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The impact of the Endangered Mauritian flying fox

Pteropus niger on commercial fruit farms and the

efficacy of mitigation
J É R É M Y S . P . F R O I D E V A U X , P A U L A . R A C E Y and G A R E T H J O N E S

Abstract The endemic Mauritian flying fox Pteropus niger is exclusively on fruits, leaves, nectar and pollen (Fleming &
perceived to be a major fruit pest. Lobbying of the Kress, ). They typically have broad diets and eat a
Government of Mauritius by fruit growers to control the fly- wide range of native, introduced and cultivated plant species
ing fox population resulted in national culls in  and , (Lobova et al., ; Fleming & Kress, ), and they can
with a further cull scheduled for . A loss of c. , in- consume large quantities of fruits (–% of their body
dividuals has been reported and the species is now categor- mass daily; Izhaki et al., ). Fruit bats are essential for
ized as Endangered on the IUCN Red List. However, until pollination and seed dispersal of at least  species of
now there were no robust data available on damage to orch- plants, of which  provide economically important re-
ards caused by bats. During October –February  we sources and products (Fujita & Tuttle, ; Lobova et al.,
monitored four major lychee Litchi chinensis and one mango ; Aziz et al., ; Vincenot et al., ). However, nega-
(Mangifera spp.) orchard, and also assessed  individual tive attitudes towards fruit bats are widespread among the
longan Dimocarpus longan trees. Bats and introduced birds general public (Vincenot et al., ; Kingston, ). Old
caused major damage to fruit, with –% fruit loss (includ- World fruit bats commonly feed on cultivated fruits, and
ing natural fall and losses from fungal damage) per tree. Bats therefore they are frequently shot, persecuted and even le-
caused more damage to taller lychee trees (.  m high) than gally culled as agricultural pests (Bumrungsri et al., ;
to smaller ones, whereas bird damage was independent of Epstein et al., ). A study in Madagascar found that
tree height. Bats damaged more fruit than birds in tall the Madagascan rousette Rousettus madagascarensis prefers
lychee trees, although this trend was reversed in small native and commercially unimportant fruit to commercially
trees. Use of nets on fruiting trees can result in as much as farmed fruit, including lychees Litchi chinensis, and main-
a -fold reduction in the damage caused by bats if nets taining supplies of native fruit may be a way of limiting
are applied correctly. There is still a need to monitor damage at orchards (Andrianaivoarivelo et al., ).
orchards over several seasons and to test non-lethal bat The Mauritian flying fox Pteropus niger is endemic to the
deterrence methods more widely. Mascarene Islands (the large islands of Mauritius, Réunion
and Rodrigues and nearby smaller islets in the south-west
Keywords Bats, Chiroptera, flying fox, fruit damage,
Indian Ocean), and in the past occurred throughout the
Mauritius, orchard, Pteropus niger, tree netting
archipelago. However, it is restricted now to the island of
Supplementary material for this article is available at Mauritius (with records of a few individuals on Réunion./S Island) as a result of habitat destruction and hunting
(Cheke & Dahl, ; Cheke & Hume, ). The species
plays a disproportionately large ecological role as a seed dis-
Introduction perser on Mauritius, especially in dispersing the seeds of
large trees that are important in forest canopies (Florens
O ld World fruit bats (Family Pteropodidae, Order
Chiroptera) are phytophagous and feed almost
et al., ).
In  the population of P. niger was estimated to com-
prise , individuals (Cheke & Dahl, ). Its range is
RYSZARD Z. OLEKSY and CHARLES L. AYADY Ecosystem Restoration Alliance Indian
Ocean, St. Pierre, Mauritius limited by habitat destruction and it is negatively affected
VIKASH TATAYAH Mauritian Wildlife Foundation, Vacoas, Mauritius by tropical cyclones, which can potentially decimate popu-
CARL JONES Durrell Wildlife Conservation Trust, Trinity, Jersey
lations (Cheke & Dahl, ; Pierson et al., ; Grant et al.,
; Cheke & Hume, ). In common with many bat spe-
JÉRÉMY S.P. FROIDEVAUX University of Bristol, Bristol, UK
cies it typically produces only a single offspring each year,
PAUL A. RACEY University of Exeter, Penryn, UK
and occasionally twins, and therefore recovery after decline
GARETH JONES (Corresponding author) School of Biological Sciences, Life is slow. The bats have been protected by law since ,
Sciences Building, University of Bristol, 24 Tyndall Avenue, Bristol BS8 1TQ,
UK. E-mail when hunting them for food and sport shooting, once
Received  May . Revision requested  July . a common practice, was made illegal. The population had
Accepted  August . increased to an estimated , by  (Republic of

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2 R. Z. Oleksy et al.

Mauritius, , ), although this estimate was based on implementation is sporadic, although it is cost-effective in
disturbance counts, which are considered to be inaccurate the long term. Government subsidies are provided for
and often result in double counting. In  the population netting in Australia, Israel and Thailand, and small net
was estimated to comprise , (± %) individuals bags are used to protect fruit on young trees in Mauritius
(Kingston et al., ). Increasingly, there are claims that and Cambodia (reviewed by Aziz et al., ). During
the bats are damaging commercial fruit crops. Mauritian – MUR  million (USD . million) was spent
fruit growers estimate that fruit bats eat , kg of lychees on the netting subsidy scheme (Agricultural Research
per annum and such damage is increasing at a rate of % and Extension Unit of the Food and Agricultural Research
annually. It is estimated that –% of fruit in orchards is Council, pers. comm.). As data on the damage caused to
damaged and, depending on market price and levels of dep- fruit crops by bats in Mauritius were contradictory and
redation, the cost of this damage can amount to MUR  not always collected systematically, we aimed to assess
million (USD . million) annually. Mauritius produces how much damage bats and other animals cause to lychee,
. , tonnes of fruit annually from , ha, with lychees mango and longan trees, and evaluated whether netting of
contributing ,–, tonnes. An estimated – trees limits the damage.
tonnes of lychees, worth MUR – million (USD .–.
million), are exported annually (Agricultural Research and
Extension Unit of the Food and Agricultural Research Study area
Council, pers. comm.; Republic of Mauritius, ).
The study was conducted in Mauritius (Fig. ), an island of
Fruit growers have lobbied the Mauritian Government
, km, which has retained , % of its high-quality native
since  (D. Sarjua, in litt. to Ministry of Agriculture, 
vegetation cover and has lost % of its native vertebrate
December , copied to Mauritian Wildlife Foundation)
species (Safford, ; Cheke & Hume, ). The remain-
to remove the legal protection for P. niger so that it may be
ing native wildlife is strongly associated with native vegeta-
controlled in orchards. In  this lobbying was successful,
tion, which has become invaded by exotic species. At least 
a cull was implemented and the official number of bats killed
plant species are highly invasive and cause degradation of
was ,, followed in  by a further cull of , bats in
native habitat. Additionally, numerous exotic animal species
an attempt to limit the damage caused to lychee and mango
cause destruction to native plants and their fruits or seeds,
farms (Republic of Mauritius, ). The cull commenced
and also spread invasive plant species (Safford, ; Cheke
despite the lack of clear evidence of the damage caused by
& Hume, ). The work was conducted during October
the Mauritian flying fox to orchard fruit (Florens, ).
–February  in four lychee orchards (Fig. ),
As a consequence of the culling, ongoing habitat loss and
Calebasses, Medine (only large, .  m, trees present),
vulnerability to cyclones, P. niger was recategorized from
Constance, and Belle Vue Maurel (only small, ,  m, trees
Vulnerable to Endangered on the  IUCN Red List,
present), and at one mango orchard in Labourdonnais.
based on an estimated population decline of % in 
Individual longan trees were assessed in St. Pierre village
and , and a projected decline of . % during –
(Moka region), as there are no longer orchards of this fruit
 (Kingston et al., ). A further cull of % of the re-
tree in Mauritius.
maining population has been announced for , and bats
continue to be hunted illegally, with an estimated ,–
, individuals killed annually (Kingston et al., ). Methods
In Mauritius the Food and Agricultural Research Council
estimated that an overall mean of % of orchard lychees Fruit assessment
were damaged annually by the Mauritian flying fox.
Ten trees were studied in each of three orchards, and the Each orchard in Mauritius typically has trees of various
mean damage to individual fruit was ,  and % (Aziz sizes, – m in height. We categorized trees as large (. 
et al., ). In contrast, a smaller study of four longan m high) or small (,  m high). Ten small and  large lychee
Dimocarpus longan trees recorded damage to all fruit pani- trees were chosen randomly and the number of fruits on
cles. Mangoes (Mangifera spp.) were also damaged at a rate each tree was assessed by counting the total number of
of –% per year (Aziz et al., ). In contrast, a study in panicles on the tree, using a tally counter, and taking the
a lychee orchard found that high winds and introduced mean number of fruits per panicle from  randomly
birds, but not bats, resulted in fruit losses of % and selected panicles. The total number of panicles was then
.%, respectively (Ramlugun, ). multiplied by the mean number of fruits per panicle. The
Netting is regarded as the only effective method for pro- same method was applied to longan trees, and mango fruits
tecting orchards from damage by frugivorous bats, and has (given their larger size) were counted individually.
been used extensively in Australia (Bicknell, ; Ullio, Randomly marked lychee trees were visited daily and the
). However, netting is expensive and therefore its number of fruits on the ground under the trees was assessed

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The Mauritian flying fox 3

Statistical analyses

Statistical analyses were conducted using R v. .. (R

Development Core Team, ). We used generalized linear
mixed models (function glmer in the lme package; Bates
et al., ) to test whether the number of fruits damaged
at two lychee orchards (Calebasses and Constance) where
both small (,  m high) and large (.  m high) trees oc-
curred (and hence bats had a choice of foraging at large
and small trees) differed between the two tree size categor-
ies. As only one small tree in the mango orchard was da-
maged by bats, confidence intervals were large and it was
not possible to run a generalized linear mixed model to in-
vestigate whether damage varied according to tree size or
fruit consumer. Given the nature of the response variable
(count data), and to handle overdispersion, we fitted the
models with negative binomial distributions. We included
the two-way interaction between tree size (large vs small)
and causes of fruit loss (bat, bird, rat, fungus, natural fall)
FIG. 1. Locations of the six orchards in Mauritius where the as fixed effects in the models, and considered the tree ID
effects of the Mauritian flying fox Pteropus niger on fruit crops nested within the orchard ID as random effects. We used
were investigated. DHARMa (Hartig, ) to build diagnostic plots and to val-
idate the model fit. We then identified the most parsimoni-
ous model amongst a set of five models that included
and counted. Based on the damage done to the fruits, they all possible variable combinations as well as the null
were categorized as bat damaged when bat canine marks model, using the second-order Akaike information criterion
(triangular shaped puncture wounds) or tooth scrape (AICc; Burnham & Anderson, ). As our best model was
marks were visible on the fruit or seed; bird damaged the most complex (interaction between the two predictors;
when the seed was smooth or the fruit had peck holes in Supplementary Table ), we conducted post-hoc contrast
the outer covering, with no visible canine punctures; fungus tests for the pairwise comparisons of interest, adjusting
infected when fruits had breaks in the skin and showed dis- P values for multiple comparisons using the Tukey method
coloration; rat damaged when marks of incisors were visible (with package lsmeans; Lenth, ).
on the fruits or seed; and natural fall when intact, undam- Bats caused more damage to large than small lychee
aged fruits were found on the ground. trees, and therefore we evaluated the effectiveness of using
It was not possible to make detailed counts of fallen fruits nets on large trees in reducing fruit loss. We conducted sep-
from longan trees because of accessibility problems and arate analyses using generalized linear mixed models with
their location near roads. Therefore, the number of empty negative binomial distributions for each orchard where the
panicles was counted three times per week and damage experiment took place (Calebasses and Medine), as different
was attributed to bats and birds combined. netting treatments were applied. The two-way interaction
between netting system (unnetted vs netted without frame
vs netted with frame for Calebasses, and unnetted vs netted
without frame only for Medine) and causes of fruit loss (bat,
Netting bird, rat, fungus, natural fall) were treated as fixed effects,
In Calebasses lychee orchard three large trees were netted and tree ID was incorporated in the models as a random
using an anti-bird net (nylon netting with a mesh of  ×  effect. We applied the same procedure for model validation
cm) placed over a frame made of PVC pipes of  mm diam- and selection as described previously (Supplementary
eter. The pipes were manipulated over the tree to lift the net Table ).
above the tree canopy. Three other trees were netted by pla-
cing the anti-bird net directly over the tree canopy without Results
frame support. The damage to the netted trees was assessed
in the same way as for unnetted trees. Medine lychee or- The numbers of trees damaged in each orchard are detailed
chard had trees already netted, with anti-bird netting placed in Supplementary Table  according to fruit type, tree size
directly over the tree canopy, with no frame support, so the and fruit consumer at unnetted trees. Lychee trees .  m
nets were in direct contact with the trees. high produced ,–, fruits on average, and trees

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4 R. Z. Oleksy et al.

,  m high produced ,–, fruits. Bell Vue Maurel

was an exception, with trees ,  m high producing only
– fruits. There was substantial variation in fruit pro-
duction among mango trees. Large trees (.  m high) pro-
duced –, fruits, whereas small trees (,  m high)
produced – fruits. Longan trees (n = ) produced
a mean of , ± SD , fruits. Overall levels of fruit loss
to all causes were substantial (–%), with three sites
having . % fruit damage (Supplementary Table ).

Bat damage

At unnetted lychee trees at Calebasses bats damaged .% of

monitored fruit on large trees and .% on small trees. At
Constance bats damaged % of fruits on large trees and
FIG. 2. Predicted means and associated % confidence intervals
.% on small trees. At Medine bats damaged .% of
of number of fruits damaged by bats and birds at large (.  m
fruit on large trees (no small trees were present). Bats da- high; n = ) and small (,  m high; n = ) trees in two lychee
maged .% of fruits on the small trees at Belle Vue Litchi chinensis orchards (Calebasses and Constance; Fig. ).
Maurel (no large trees present). At mango trees bats da- Model predictions arise from the most parsimonious generalized
maged .% of fruits on large trees and .% of fruits on linear mixed model that included the interaction between the
small trees. Bat damage was therefore more severe at large cause of damage (bat, bird, fungus, natural fall, rat) and tree size
trees compared with smaller ones for both lychees and (large, small). P values adjusted for multiple comparisons are
given for the pairwise comparisons of interest. The full
mangos (although statistical analysis was not possible for
description of these comparisons is in Supplementary Table .
the latter). This height-dependent effect was significant
for bat damage at lychee orchards, but damage by all
other agents was not related to tree height (Fig. ; accounted for .% of total fruits produced by the large
Supplementary Table ). mango trees and .% at the small mango trees studied.

Bird damage Effects of netting

Birds damaged .% of fruit on large lychee trees at The effects of netting were assessed at two lychee orchards,
Calebasses, and .% on small trees. The equivalent values Calebasses and Medine (Supplementary Table ). At
at Constance were . and .%, respectively. For the Calebasses bats damaged only .% of fruits at large trees
large lychee trees at Medine, .% of fruits were damaged netted over a frame (n = ), whereas birds damaged .%
by birds. For the small lychee trees at Belle Vue Maurel, of fruits, compared with .% bat damage and .% bird
.% of fruits sustained bird damage. When large and damage at unnetted control sites. The total loss of fruits
small lychee trees were present, bird damage was signifi- was .% for trees netted with frames, compared with
cantly lower than bat damage at large trees, although birds .% at unprotected trees. At the trees netted with no
caused more damage than bats at smaller trees (Fig. ; frame (n = ) bats accounted for .% of damage to fruits,
Supplementary Table ). and birds for .%. The total loss was .% on netted
At mango trees birds damaged . and .% of fruits on trees with no frame, compared with .% on unprotected
large and small trees, respectively. Damage by birds at small trees. Model predictions (Fig. ; Supplementary Table ) in-
mango trees was therefore substantial and considerably dicated that netting both with and without frames at
higher than that caused by bats. For longan trees, .% of Calebasses significantly reduced fruit damage by bats but
fruits were damaged by either birds and/or bats. had no effect on bird damage. There was no significant dif-
ference in damage by bats according to whether the nets
were on frames or not.
Other damage and natural fall At trees netted without frames at Medine the damage
caused by bats decreased to .%, compared with .% at
Damage by rats was negligible (, % of all fruits). Loss by unnetted trees (Supplementary Table ). Birds accounted for
fungal infection was generally small, although it reached .% of damage at netted trees without frames, compared
–% on lychee trees at Constance. Fruit loss by natural with .% at unnetted trees at Medine. Netting trees without
fall was typically , % for lychee trees, although it frames had no significant effect on bird damage but had a

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The Mauritian flying fox 5

FIG. 3. Predicted means and

associated % confidence
intervals of number of fruits
damaged by bats and birds at
large trees in one lychee orchard
(Calebasses; Fig. ) in relation to
the netting system implemented:
unnetted (n = ), netted without
frame (n = ), netted with frame
(n = ). Model predictions arise
from the most parsimonious
generalized linear mixed model
that included the interaction
between the cause of damage
(bat, bird, fungus, natural fall,
rat) and the netting system. P
values adjusted for multiple
comparisons are given for the
pairwise comparisons of interest.
The full description of these
comparisons is in Supplementary
Table .

large and significant effect on reducing bat damage (Fig. ;

Supplementary Table ). The total damage to fruits at
Medine was .% on trees netted without a frame, com-
pared with .% on unnetted trees.


The damage caused by bats and other animals varies from

orchard to orchard. However, small trees (,  m) were
less prone to damage by bats compared to large trees (. 
m). Across the four lychee orchards studied, bats damaged
.% of fruits overall and .% of fruits at small trees, on
average, whereas birds damaged .% of fruits overall and
.% of fruits at small trees. Bat damage was measured at
the trees that we monitored, but bats can potentially carry
FIG. 4. Predicted means and associated % confidence intervals
fruit away to consume it elsewhere. However, from our ob-
of number of fruits damaged by bats and birds at large trees in
servations we believe that in general the bats ate fruit at the one lychee orchard (Medine; Fig. ) in relation to the netting
trees from which they took it, and therefore we believe that system implemented: unnetted (n = ), netted without frame
our estimates of damage are realistic. (n = ). Model predictions arise from the most parsimonious
Pruning trees could minimize the impact bats have on generalized linear mixed model that included the interaction
fruit crops. However, in places such as Medine and between the cause of damage (bat, bird, fungus, natural fall, rat)
Constance the damage caused by bats was pronounced and the netting system. P values adjusted for multiple
comparisons are given for the pairwise comparisons of interest.
and even small trees in those orchards suffer substantial
The full description of these comparisons is in Supplementary
fruit losses. Flying foxes often prefer to eat fruits that are Table .
too ripe to be sold (Singaravelan et al., ), and therefore
harvesting the fruit at an earlier stage could also reduce the
attractiveness of orchards to bats and other frugivores (Aziz Birds such as the invasive exotic red-whiskered bulbul
et al., ). Up to % of lychee loss in this study was as a Pycnonotus jocosus and common myna Acridotheres tristis,
result of natural falling of fruit. This loss was even higher at and to a lesser extent the ring-necked parakeet Psittacula
the mango orchard, exceeding % of total fruit at large krameri cause considerably more damage than bats to smal-
trees. These losses could potentially be reduced by earlier ler trees across all the orchards (mango and lychee). Other
harvesting. animals, including rats, were found to have little to no

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6 R. Z. Oleksy et al.

impact on the fruits. This could be because rats may feed on are pruned low because it makes them easier to harvest, a
fruits that are already on the ground. Fungal infection of ly- practice that should be applied to all mango and lychee
chee fruits is common in Mauritius. However, in all of the trees. Plastic sheets are also used to protect longan orchards
orchards assessed, fungicide was sprayed on the trees to pro- in Thailand, and may also accelerate fruit ripening (Aziz
tect the fruits against such infection, and loss to fungal in- et al., ). It is likely that large-scale fruit growers in
fection was minimal. Mauritius may benefit from using netting in orchards
Netting was successful in reducing the damage caused by even though they do not receive government subsidies,
bats –-fold in Calebasses and more than -fold in given the substantial reductions in fruit damage resulting
Medine, compared with unnetted trees. It is therefore im- from netting, and the relatively low cost of nets.
perative to net trees, preferably using frames so that the Currently, apart from netting, growers in Mauritius use a
net is held above the tree canopy. If nets are placed directly range of other methods, including burning leaves or tyres to
on the trees and in contact with fruits, bats can land on and create dense smoke, which the bats avoid; installing light
feed through the net. Additionally, black nets placed directly bulbs on every tree to illuminate the area and make it less
over the trees are less visible to the bats and as a result bats attractive to the bats; and using plastic flags placed above
occasionally becoming entangled in them. To avoid entan- the tree canopy, firecrackers and shotguns to frighten the
gling of bats and fruit damage through the net, it is advised bats, and in the latter case sometimes to kill them.
to make the net as conspicuous as possible against the tree. Additionally, a dead bat may be attached to a bamboo
Use of white rather than black nets is recommended. pole and placed over the tree canopy to deter other bats.
Bird damage on netted trees increased compared with None of these methods is known to be particularly effective
unnetted trees, possibly because birds may become trapped in reducing the impact of bats on fruit, but fruit growers pre-
after entering the canopy through gaps in the net. They fer them to netting because they are less time- and labour-
cause more damage to the fruits in such cases because intensive. Netting is not well accepted by farmers in
they continue feeding on them. To avoid gaps, wider nets Mauritius, and it is impractical to net tall trees (.  m
($  m) than those commonly available in Mauritius high), which are common in backyards.
(which come in rolls  ×  m) are desirable so that the In Madagascar, Plantskydd deer repellent (Tree World
whole tree can be covered without the need to join nets to- Plant Care Products, Inc., Sechelt, Canada) was tested on ly-
gether, as is the typical practice in Thailand (Aziz et al., chee trees as a non-lethal taste deterrent against fruit bats
). Alternatively, birds may fly under the nets if they (Raharimihaja et al., ). The use of this deterrent was
are not secured around the base. The ideal is to have fruit compared to waving of red flags and ringing bells.
trees entirely caged to exclude bats and birds. Plantskydd proved to be successful in mitigating bat damage
In  the Mauritian Government subsidized % of the to lychees. However, its composition of dried blood makes
cost of five nets ( ×  m) per backyard grower, and % of its use inappropriate in Mauritius because of religious be-
netting costs for orchards , . ha in area (Republic of liefs and veterinary health restrictions on its import.
Mauritius, ). Nets typically cost MUR ,–, (c. However, alternative repellents that are not blood-based
USD –), and are reusable across years if they are stored could be tested. Methods using decoy crops, light or
appropriately. Small-scale fruit growers can potentially re- sound, and advancing fruit harvest time may have potential
coup the investment in nets, and potentially even the full for reducing fruit loss to bats (Aziz et al., ).
cost of the nets within the first year of use. The success of This work has shown the extent of fruit loss in Mauritian
the scheme is demonstrable and it has been renewed annu- orchards as a result of bird and bat predation and other fac-
ally since . However, major orchards in Mauritius may tors, and is a first step in documenting damage and suggest-
have . , fruiting trees, and therefore the subsidy only ing solutions. It is important to monitor the orchards for
meets the needs of small-scale growers. In Australia, more seasons, to establish the pattern of fruit loss over
where orchards can be as large as  ha (Aziz et al., ), time. In years when natural food is scarce, bats may be
full exclusion netting is used, which gives % protection forced to feed more extensively on commercial fruits,
against bats, birds and abiotic factors. The netting has a whereas during times when native fruits are available the
mesh size of c.  mm, is erected well above the height of damage may be less pronounced.
the trees and is attached to the ground on the sides. A simi- In conclusion, we are optimistic that netting fruit trees
lar approach is used in Israel on mango, lychee and banana would reduce the loss of commercial fruit to birds and
plantations. In many cases the initial reason for using exclu- bats considerably if installed appropriately. The Mauritian
sion netting was to reduce water evaporation, as irrigation in Wildlife Foundation continues to promote the benefits of
Israel is expensive; however, netting ultimately protects the netting to the Government of Mauritius, which reviews
crops against fruit bats and birds. Additionally, the trees are and changes its Fruit Protection Scheme regularly, and a
kept low and trimmed regularly, which makes netting easier strong evidence base exists showing the causes of fruit loss
(authors, pers. obs.). In some orchards in Mauritius, trees at orchards, and how netting can reduce these losses

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The Mauritian flying fox 7

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