ArticlePDF Available
Afr J Ecol. 2020;00:1–4.
  1© 2020 John Wiley & Sons Ltd
Received: 8 Octob er 2019 
  Revised: 26 Ap ril 2020 
  Accepted: 27 April 2020
DOI: 10.1111/aje.12749
Seawater ingestion by the Mauritius flying fox
Vashist Omprasad Seegobin1| Jean-Michel Probst1,2
1Depar tment of Bioscie nces and Ocean Stu dies, Tropi cal Island Biod iversi ty, Ecolog y and Conservation Pole of Research, University of Mauritius , Le Rédui t,
2Association Nature & Pat rimoine, Réuni on, Réun ion
Correspondence: Vashist Omprasad Seegob in, Tropica l Island Biodive rsity, Ecology and Conse rvation Pole of Re search, Depa rtme nt of Biosciences a nd Ocean
Studies , Univer sity of M auritius, Le Réduit, Mauriti us.
Mauritius (centred around 20° 20’ S; 57° 34’ E; area 1,865 km2;
828 m maximum elevation) is a 7.8-million-year old volcanic island
with moist-to-wet tropical climate, located around 900 km east of
Madagascar in the Indian Ocean. Anthropogenic impacts started
shortly before human colonisation in 1638, which itself led to the
island becoming one of the most ecologically devastated worldwide
(Cheke & Hume, 2008; Florens, 2013; Hammond et al., 2015).
Flying foxes face threats mainly from hunting, natural habitat
transformation, invasive alien species and climate change (Vincenot,
Florens, & Kingston, 2017) which is likely to increase the occurrence
and intensit y of stochastic events such as cyclones (Elsner, Kossin, &
Jagger, 2008; Kishtawal, Jaiswal, Singh, & Niyogi, 2012; Kuleshov, Qi,
Fawcett, & Jones, 2008; Webster, Holland, Curry, & Chang, 2005).
In Mauritius, invasive alien species largely contribute to declining
native foraging habitat quality of the ‘Endangered’ Mauritius flying
fox (Pteropus niger) (Cheke & Hume, 2008; Krivek, Florens, Baider,
Seegobin, & Haugaasen, 2020). Furthermore, cyclones accompanied
with heavy rainfalls and gusts (around 100 km/hr) previously caused
60% cultivated fruit loss in Mauritius (Anon., 2012, 2015), potentially
exacerbating fruit availability for species like P. niger (McConkey,
Drake, Franklin, & Tonga, 2004; Soto-Centeno & Kurta, 2006).
Other flying fox species in Comoros, Papua New Guinea and
Philippines are known to skim over the sea before licking their
fur to obtain minerals absent from their frugivorous diet (Iudica &
Bonaccorso, 2003; Probst & Winter, 1993; Stier, 2003). However,
Thomas (1984) suggested that obligate frugivorous bats adjust nutri-
ent demands by varying amounts of eaten fruits. In Australia, Pteropus
poliocephalus frequently body dips in freshwater during days of high
temperature for cooling and rehydration (Rakotopare & Abhaya, 2019).
Eight afternoon kayak expeditions were conducted over three
weeks between February and beginning of April 2018 in the lagoon
of Pointe d’Esny, Blue Bay and from the beach (located at 20° 26'
41.56" S; 57° 42' 21.25" E). More obser vations were ma de in the bay
of Mahebourg (20° 24' 50.54" S; 57° 42' 40.60" E) and La Cambuse
(20° 27' 26.67" S; 57° 41' 38.04" E) in Februar y 2019. A Nikon D3200
DSLR camera and Sigma 100 – 300 mm telephoto lens were used to
photograph observations.
The first observ ation was made in Februar y 2 018 in the lago on of
Pointe d’Esny at around 200 m from the coast. A Mauritius flying
fox slowly approached the water surface, within 50 m from our
position, and after several attempts clumsily succeeded in dip-
ping its lower belly part into the water and immediately licked
its wet fur before flying towards the mainland (Figure 1). Three
similar observations were made the next day and fifteen in
total over eleven days. Most observations occurred within five
weeks following heavy rainfall in Januar y (794 mm) (Statistics
Mauritius, 2018) and cyclonic conditions with gusts of 120 km/
hr (reliefweb, 2018) and after another cyclone in mid-February
2019. Each observation lasted around fifteen seconds, sometimes
occ ur ri ng on cloudy and rainy days, be tween Februar y and b egin-
ning of April, during temperature averaging 25°C, minimal wind
and calm sea conditions, and between 15:00 and 19:00. Similar
dipping behaviour throughout the year remains possible although
not reported.
Frugivorous species commonly lack certain minerals like sodium
and iron (Stier, 2003; Studier, Sevick, Ridley, & Wilson, 1994) and
often compensate through salt licking (Iudica & Bonaccorso, 2003;
Klaus & Schmidg, 1998). However, environmental factors such as
strong cyclonic winds, rainfall and temperature may influence avail-
ability (Grant, Craig, & Trail, 1997; Karr, 1976; Remis, 1997) and
quality of fruits (Sams, 1999; Worman & Chapman, 2005), alter-
ing feeding habits (Elangovan, Marimuthu, & Kunz, 1999; Kunz &
Fenton, 2005;Raghuram, Singaravelan, Nathan, & Emmanuvel, 2011).
Stochastic events like cyclones and torrential rainfall (794 mm in
January 2018 compared to 146 and 185 mm for January 2017 and
2016, respectively (Statistics Mauritius, 2018)), potentially reduce
foraging possibilities of Pteropus niger (Cheke & Dahl, 1981), beyond
consequences emanating from native foraging habitat destruction
and degradation from alien species invasion (Florens & Baider, 2013;
Florens et al., 2016; Florens, Baider, Seegoolam, Zmanay, &
Strasberg, 2017). Furthermore, invasive long-tailed macaques pop-
ulation (Macaca fascicularis) increased markedly in the last dozen of
years and the species is known to consume large numbers of na-
tive fruits, thereby depleting fruit resources otherwise available for
P. niger (Laurance & Peres , 2006). Such situation could prompt flying
foxes to explore alternative ways to supplement their diet. However,
food scarcity as a driver for seawater ingestion remains uncertain.
Se a wat e r coul d pro v ide a ri ch so u r ce of so d ium an d chl ori d e min-
erals for flying foxes (Stier, 2003). Despite fruit availability, dipping
behaviour by Pteropus niger might hel p ob tain nu trients that is sc arce
in consumed fruits. Similar dipping behaviour has been observed in
other flying fox species in the Comoros islands by P. seychellensis co-
morensis (Probst & Winter, 1993;Stobb, 1994), in the Philippines by
P. vampyrus and Acerodon jubatus (Stier, 2003), in the coas tal regions
of Papua New Guinea by P. conspicillatus, P. hypomelanus and P. t on-
ganus (Iudica & Bonaccorso, 20 03) and sug ge st ed to be either for ac-
quiring nutrient s, thermoregulation or protecting against parasites.
Other flying fox species were commonly obser ved body dipping
in freshwater during hot periods for cooling and rehydration. Pteropus
poliocephalus and P. a le c to have been seen licking their fur after
freshwater body dipping (Markus & Blackshaw, 2002; Rakotopare
& Abhaya, 2019). Commerson (Buffon, 1783) observed P. niger, in
Reunion, to occasionally dip their bodies in fresh water. However,
these behaviours, triggered by warm temperatures, seemed differ-
ent from our observations whereby dipping occurred in sea water,
late afternoons with temperature averaging 25°C, sometimes on
cloudy and rainy days.
To our knowledge, this is the first record of seawater ingestion by
Pteropus niger. Some local people and fishermen who witnessed this
behaviour assumed that the species was fish-hunting, causing re-
sentment that could favour its persecution. Our observations can
help reduce incorrect beliefs and improve underst anding of the
feeding behaviour versatility, resilience ability and local adapta-
tion of P. niger to particular opportunities within its environment.
Additionally, seawater body dipping to protect flying foxes from
parasites could potentially be explored.
The authors declared that they do not have any potential sources of
conflict of interests.
FIGURE 1 (a) A Mauritius flying fox
approaching the seawater sur face, (b)
dipping it s lower body part into it and (c)
licking the salty water from its wet fur
before (d) flying back to the mainland
(a) (b)
(c) (d)
The data that support the findings of this study are openly available
in ‘Dryad’ at w3rc (Seegobin,
Vashist Omprasad Seegobin https://orcid.
Anon. (2012). Mauritius: Tropical cyclone Dumile has severely affected
the fruit sector. Retrieved Februar y 23 2020, from https://en.indi- tius-Tropi cal-cyclo ne-Dumile-has-sever
ely-affec ted-the-fruit-sector_a692.html.
Anon, (2015). Mauritius: Vegetable and fruit crops severely impacted after
the passage of tropical cyclone Bansi. Retrieved February 22 2020,
from tius-Veget able-and-
fruit-crops-sever ely-impac ted-after-the-passa ge-of-tropi cal-cyclo
Buffon, G. L. (1783). Histoire naturelle des animaux. Tome 1-10. années
1770-1783. Paris
Cheke, A., & Dahl, J. (1981). The status of bats on western Indian Ocean
islands, with special reference to Pteropus. Mammalia, 45(2), 205–
Cheke, A., & Hume, J. (2008). Lost Land of the Dodo. An Ecological History
of Mauritius, Réunion & Rodrigues. London: T & AD Poyser.
Elangovan , V., Marimuthu, G., & Kunz, T. (1999). Temporal patterns of
individual and group foraging behaviour in the shor t-nosed fruit bat,
Cynopterus sphinx, in south India. Journal of Tropical Ecology, 15(5),
Elsner, J. B., Kossin, J. P., & Jagger, T. H. (2008). The increasing intensity
of the strongest tropical cyclones. Nature, 455(7209), 92–95.
Florens, F. B. V. (2013). Conservation in Mauritius and Rodrigues: Challenges
and achievements from two ecologically devastated oceanic islands.
Conser vation Biolog y: Voices from the tropics, 40–50.
Florens, F. B . V., & Baider, C. (2013). Ecological restoration in a devel-
oping island nation: How usef ul is the science? Restoration Ecology,
21(1), 1–5. /10.1111/j.1526-100X.2012.00920.x
Florens, F. B. V., Baider, C., Martin, G. M., Seegoolam, N. B., Zmanay, Z., &
Strasberg, D. (2016). Invasive alien plants progress to dominate pro-
tected and best-preserved wet forest s of an oceanic island. Journal
for Nature Conservation, 34, 93–100. htt ps://
Florens, F. B. V., Baider, C., Seegoolam, N. B., Zmanay, Z., & Strasberg,
D. (2017). Long-term declines of native trees in an oceanic island's
tropical forests invaded by alien plant s. Applied Vegetation Science,
20(1), 94–105.
Grant, G. S., Craig, P., & Trail, P. (1997). Cyclone-induced shift in foraging be-
havior in flying foxes in American Samoa. Biotropica, 29(2), 224–228.
Hammond, D. S., Gond, V., Baider, C., Florens, F. B. V., Persand, S., &
Laurance, S. (2015). Threats to environmentally sensitive areas from
peri-urban expansion in Mauritius. Environmental Conservation, 42(3),
256–267. 89291 4000411
Iudica , C., & Bonaccorso, F. (2003). Anecdotal observations of seawa-
ter ingestion by flying foxes of the genus Pteropus (Chiroptera:
Pteropodidae). Mammalia, 67(3), 455–458.
Kar r, J. R. (1976). Seasonality, re source availability, an d com munit y div er-
sity in tropical bird communities. The American Naturalist, 110 (976),
973–994. https://doi.or g/10.1086/283121
Ki sht aw al, C., Jai swa l, N., Sing h , R. , & Ni yo gi, D. (2012). Trop ica l cyclo n e inte n-
sification trends during satellite era (1986–2010). Geophysical Research
Letter s, 39(10), L10810. L05170 0
Klaus, G., & Schmidg, B. (1998). Geophag y at natural licks and mam-
mal ecology: A review. Mammalia, 62(4), 482–498. https://doi.
Krivek, G., Florens, F. B. V., Baider, C., Seegobin, V. O., & Haugaasen, T.
(2020). Invasive alien plant control improves foraging habitat quality
of a threatened island flying fox . Journal for Nature Conservation, 54,
Kuleshov, Y., Qi, L., Fawcett, R., & Jones, D. (2008). On tropical cyclone
activity in the Southern Hemisphere: Trends and the ENSO con-
nection. Geophysical Research Letters, 35(14), L14S0 8. ht tp s://doi.
org /10.1029/2007G L032983
Kunz, T. H., & Fenton, M. B. (2005). Bat ecology: University of Chicago
Laurance, W. F., Peres, C. A. (20 06). Emerging threats to tropical forests
(pp. 199–214). Chicago, IL: Universit y of Chicago Press.
Markus, N., & Blackshaw, J. K. (2002). Behaviour of the black flying fox
Pteropus alecto: 1. A n ethogram of behaviour, and preliminary char-
acterisation of mother-infant interactions. Acta Chiropterologica, 4(2),
13 7–1 5 3 .
McConkey, K . R., Drake, D. R., Franklin, J., & Tonga, F. (20 04). Effects
of Cyclone Waka on flying foxes (Pteropus tonganus) in the Vava'u
Islands of Tonga . Journal of Tropical Ecology, 20(5), 555–561.
Probst , J. M., & Winter, M. (1993). Trempage claqué aux ras des vagues :
Pteropus seychellensis Milne-Edwards, 1877 (Roussette des Seychelles).
Comores, Mayotte. Observation Mascarines, 6, 89.
Raghuram, H., Singaravelan , N., Nathan, P. T., & Emmanuvel, K. (2011).
Foraging ecology of Pteropodid bat s: Pollination and seed dis-
persal. In J. Zupan, & S . Mlakar (Eds.), Bats: Biology, Behaviour and
Conservation (pp. 177–188). New York, USA : Nova Science.
Rakotopare, N., & Abhaya, K. (2019). Comportement de trempage:
Pteropus poliocephalus Temminck, 1825 (Roussette à tête grise).
Australie. Données Naturalistes Animalières, 23, 72.
reliefweb, , (2018). Mauritius, Tropical Storm BERGUITTA Emergency Plan
of Action (EPoA) DREF n° MDRMU002. Retrieve d February 20 2020,
from https://relie t/mauri tius/mauri tius-tropi cal-
storm-bergu itta-emerg ency-plan-action-epoa-dref-n-mdrmu002.
Remis, M. J. (1997). Western lowland gorillas (Gorilla gorilla go-
rilla) as seasonal frugivores: Use of variable resources. American
Journal of Primatology, 43(2), 87–109. ht tps://doi.o rg/10.1002/
Sams, C. E. (1999). Prehar vest fac tors affecting postharvest texture.
Postharvest Biolog y and Technology, 15(3), 249–254. https://doi.
org /10.1016/S 0925-5214(98) 00 098-2
Seegobin, V. (2019). Seawater body dipping behaviour of the Mauritian fly-
ing fox. Retrieved from: w3rc.
Soto-Centeno, J. A., & Kurt a, A . (20 06). Diet of t wo nectarivorous
bats, Erophylla sezekorni and Monophyllus redmani (Phyllostomidae),
on Puer to Rico. Journal of Mammalogy, 87(1), 19–26. https://doi.
org /10.16 44/05-MAM M-041R1 .1
Statistics Mauritius. (2018). Republic of Mauritius: Mauritius in Figures
2016. Ministry of Finance and Economic Development. Retrieved
February 21 2020, from http://stats mauri sh/
P u b l i c a t i o n s / P a g e s / M a u r i t i u s - i n - F i g u r e s . a s p x .
Stier, S. C. (2003). Dietary habits of two threatened co-roosting flying foxes
(Megachiroptera) Subic Bay Philippines. (MSc), The University of Montana.
Stobb, R. (1994). Piscivory in the Comoro Islands flying fox Pteropus seychellen-
sis comorensis-a refutation. South Afr ican Journal of Sc ience, 90(5), 264–265.
Studier, E. H., Sevick, S. H., Ridley, D. M., & Wilson, D. E. (1994). Mineral
and nitrogen concentrations in feces of some neotropical bats. Journal
of Mammalogy, 75(3), 674–680.
Thomas, D. (1984). Fruit intake and energy budget s of frugivorous bats.
Physiological Zoology, 57(4), 457–467.
Vincenot, C. E., Florens, F. B. V., & Kingston, T. (2017). C an we protect
island flying foxes? Science, 355(6332), 1368–1370.
Webster, P. J., Holland, G. J., Curry, J. A., & Chang, H.-R . (2005). Changes
in tropica l cy cl on e nu mber, dura ti on , an d in te ns it y in a warmi ng envi-
ronment. Science, 309(5742), 184 4–1846.
Worman, C. O. D., & Chapman, C. A . (2005). Seasonal variation in the
qualit y of a tropical ripe fruit and the response of three frugivores.
Journal of Tropical Ecology, 21(6), 689–697. ht tps://doi.or g/10 .1017/
S 0 2 6 6 4 6 7 4 0 5 0 0 2 7 2 5
How to cite this article: Seegobin VO, Probst J-M. Seawater
ingestion by the Mauritius flying fox. Afr J Ecol. 2020;00:1–4.
https://doi .org /10.1111/aje.12749
Full-text available
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Pteropodid bats are important aerial pollinators and seed dispersers in the Old World tropics and thus play a key role in forest dynamics and regeneration. In India, among the 13 species of pteropodid bats, the foraging ecology of four species Cynopterus sphinx, C. brachyotis, Rousettus leschenaulti and Pteropus giganteus were extensively studied. These species occur sympatrically and forage on resources like leaves, flowers and fruits, and inadvertently effect pollination and seed dispersal of the resource trees. These bats either consume fruits and flowers in situ or carry them to nearby roosts for feeding ex situ depend on the carrying capacity of bats and size and the nutritional quality of consuming resource. We suggest that small pteropodid bats aid short distance pollination and the dispersal of seeds of small sized fruits, whereas large pteropodid bats aid long distance pollination and the dispersal of seeds of large sized fruits with some exceptions that depended on the mass of the fruit. At least 300 plant species of nearly 200 genera rely mainly on these fruit bats for their propagation. Some of the plants are economically important and they produce approximately 500 economically valuable products including fruits, dyes, tannins, timber, medicines, fiber and fire wood. In this review, we discuss the foraging ecology of pterpodid fruit bats in the Old world tropics particularly with reference to chiropterophilly and zoochory by fruit bats.
Flying foxes provide critical ecosystem services by pollinating and disseminating diverse plant species. Yet, they face intensifying threats, particularly on islands. The situation is epitomized by the Indian Ocean island of Mauritius. In December 2016, the Mauritian government implemented the second mass cull of a threatened, endemic flying fox species, Pteropus niger, in 2 years. Government figures suggest that at least 45% of the overall P. niger population of just over 90,000 individuals were culled during the two campaigns; illegal killing and incidental mortality of pups during the culls raise likely losses to over 50%. The dire situation of island flying foxes worldwide calls for effective, science-based conservation strategies to prevent further loss of biodiversity and function.
The Mascarene islands in the southern Indian Ocean - Mauritius, Réunion and Rodrigues - were once home to an extraordinary range of birds and reptiles. Evolving on these isolated volcanic islands in the absence of mammalian predators or competitors, the land was dominated by giant tortoises, parrots, skinks and geckos, burrowing boas, flightless rails & herons, and of course (in Mauritius) the Dodo. Uninhabited and only discovered in the 1500s, colonisation by European settlers in the 1600s led to dramatic changes in the ecology of the islands; the birds and tortoises were slaughtered indiscriminately while introduced rats, cats, pigs and monkeys destroyed their eggs, the once-extensive forests logged, and invasive introduced plants from all over the tropics devastated the ecosystem. The now-familiar icon of extinction, the Dodo, was gone from Mauritius within 50 years of human settlement, and over the next 150 years many of the Mascarenes' other native vertebrates followed suit. The product of over 30 years research by Anthony Cheke, Lost Land of the Dodo provides a comprehensive yet hugely enjoyable account of the story of the islands' changing ecology, interspersed with human stories, the islands' biogeographical anomalies, and much else. Many French publications, old and new, especially for Réunion, are discussed and referenced in English for the first time. The book is richly illustrated with maps and contemporary illustrations of the animals and their environment, many of which have rarely been reprinted before. Illustrated box texts look in detail at each extinct vertebrate species, while Julian Hume's superb colour plates bring many of the extinct birds to life. Lost Land of the Dodo provides the definitive account of this tragic yet remarkable fauna, and is a must-read for anyone interested in islands, their ecology and the history of our relationship with the world around us. - See more at:
How did the native and alien woody plant communities of protected lowland wet forests of a tropical oceanic island change in the presence of understorey invasive alien plants over the medium (21–27 yrs) and longer term (68 yrs)? Bel Ombre, Brise Fer and Macchabé forests in the Black River Gorges National Park (BRGNP), Mauritius (20°22′10″–20°28′17″ S, 57°24′45″–57°27′12″ E). Random and replicated vegetation plots were sampled in two protected forest areas whose communities of woody plants were surveyed 21 and 27 yrs previously, and in another protected and similar forest that was surveyed 68 yrs earlier, to identify species and measure stem diameters so as to permit comparisons at the three sites over these time periods. Invasion by woody alien plants has progressed through time at all three sites, comprised mostly of the understorey and shade-tolerant Strawberry guava (Psidium cattleianum). Concomitantly, although reductions in native woody plant species richness of large trees (≥10 cm DBH) were not statistically significant at the community level, their densities and basal area had roughly halved in 68 yrs. Comparisons with the studies 21–27 yrs previously confirmed the tendency towards a reduction in density of larger trees, besides indicating a trend towards decline in species richness of smaller understorey trees. An unabated replacement of native by alien trees over the longer term is occurring in protected areas of the lowland wet forests of the BRGNP and presumably in other similarly invaded areas. Our work stresses the importance of long-term monitoring in elucidating impacts of invasive alien plants at the community level, particularly when dealing with slow-growing and long-lived species. Plausible mechanisms bringing about the observed changes and that have been found at the same or similar sites, include reduced regeneration, growth, production of flowers and fruits and increased mortality of native trees when these grow among invasive alien plants. The control of invasive alien plants, where they invade tropical forests, should be a central conservation objective even in better preserved and well protected areas.
Concentrations of nitrogen and minerals in individual fecal pellets of Noctilio leporinus directly reflect composition of the fish, crustaceans, beetles, or moths originally consumed. Among Neotropical bats that feed primarily on plant parts (fruits, nectar, pollen, and flowers), animal parts (insects and vertebrates), or both, differences in measured concentrations of nutrients are present in feces. Nitrogen levels are markedly higher and sodium levels are marginally higher in feces of carnivores and omnivores than in frugivores. Calcium levels are higher and potassium levels are lower in feces of bats that primarily consume insects. Total iron levels in feces of frugivorous species are marginally lower than in carnivores or omnivores. Magnesium concentrations seem unrelated to feeding habits. Intake of nitrogen, potassium, and magnesium appears to be adequate for bats of all feeding habits. Periodic deficiencies for calcium exist for insectivorous species and for sodium, and possibly iron, in some frugivorous species.