TY - JOUR
AU - Jones, Gareth
AB - Abstract Bats provide important pollination and seed-dispersal services to native angiosperms. However, many bat species are increasingly threatened by human disturbance, including the Mauritian flying fox (Pteropus niger), an endemic, keystone seed disperser. Native forests are scarce and P. niger frequently feeds in commercial plantations, where it now is considered a pest and subjected to frequent culling, thereby hindering conservation efforts. The invasive long-tailed macaque (Primates: Cercopithecidae, Macaca fascicularis) potentially competes with P. niger for scarce native fruits. We investigated the extent of dietary overlap between M. fascicularis and P. niger on Mauritius by sampling fruit drop for 17 tree species and identifying additional food species along line transects. Fruits of 13 of 17 species were eaten by animals and fruit production across tree replicates generally was low but highly variable. Although M. fascicularis ate only 4% of fruit overall, they consumed 20–100% of the fruits of seven species. Approximately 39% of dropped fruits were intact; based on field observations, most probably were dropped by M. fascicularis. Unlike P. niger, M. fascicularis ate mostly unripe fruit and depleted all fruit of certain species at an unripe stage. Hence, M. fascicularis may restrict P. niger’s diet and potentially disrupt seed dispersal of some tree species. Furthermore, small trees are more prone to fruit depletion at an unripe stage by macaques. In addition, asynchronous fruiting phenology across forest fragments may modulate the provision of native fruits to P. niger throughout the year. Although competition can be demonstrated only by controlled experimental studies that are logistically impossible in our scenario, our results highlight potential detrimental consequences that introduced frugivores may have on keystone seed dispersers. Finally, our results suggest that a more integrative and island-wide approach to forest restoration may be valuable for the conservation of P. niger. feeding ecology, introduced primates, invasive alien species, oceanic island, Pteropus niger Bats form a major part of mammalian diversity and ~25% of bat species are confined to islands (Jones et al. 2009), including most Pteropus species (flying foxes, Pteropodidae) (Rainey and Pierson 1992). Flying foxes mainly are frugivores or nectarivores (Banack 1998) and act as seed dispersers and pollinators for many plants (Cox et al. 1991; Nyhagen et al. 2005; McConkey and Drake 2015; Aziz et al. 2017) as they are capable of long-distance dispersal (Richards 1990; Oleksy et al. 2017), have an acute sense of smell that allows them to distinguish ripe from unripe fruit (Hodgkison et al. 2003), and can enhance seed survival and germination through pulp removal and ingestion (Traveset 1998; Oleksy et al. 2017; Saldaña-Vázquez et al. 2019). Furthermore, flying foxes can encourage the regeneration of degraded tropical forests by maintaining community diversity and promoting gene flow of dominant forest trees (Banack 1998). Island archipelagos have become hotspots for threatened bat species (Frick et al. 2019) and island-dwelling bats, including Pteropus spp., are increasingly threatened by human disturbance (Mickleburgh et al. 2002; Rocha 2015; Vincenot et al. 2015, 2017). Invasive alien species (IAS) have caused damage to oceanic island ecosystems globally, leading to unprecedented rates of species extinction (Paulay 1994; Tye et al. 2018). Oceanic islands usually exhibit disproportionate levels of species richness and endemism (Kueffer and Kinney 2017), and island biotas are particularly vulnerable to the impacts of IAS (Bellard et al. 2016; Groombridge et al. 2018). As a result, islands often exhibit high levels of endangerment at several taxonomic levels (Pimm et al. 2014), and most bats threatened by IAS inhabit islands (Welch and Leppanen 2017). IAS can disrupt pollination and seed dispersal by fruit bats (Traveset and Richardson 2006), which could have deleterious consequences for island ecosystems. Many islands have lost large proportions of their frugivore communities, including large-bodied seed dispersers (Hansen and Galetti 2009; Heinen et al. 2018). This loss has reduced seed-dispersal functions that are key to plant regeneration (Redford 1992; Hansen and Galetti 2009; Wright et al. 2009; Heinen et al. 2018). On some islands, large Pteropus species are regarded as the last keystone dispersers for large-seeded plants because of the absence of other large frugivores (Cox et al. 1991; McConkey and Drake 2002; Florens et al. 2017a). However, threats of IAS to bats and their ecological roles remain poorly assessed (Frick et al. 2019). Mauritius exemplifies how IAS may degrade forest ecosystems and affect frugivore communities (Cheke and Hume 2008). Rats (Rattus rattus), feral cats (Felis catus), and Chinese guava (Psidium cattleyanum) are among the many invasive species implicated both in extinctions (Cheke and Hume 2008; Cheke 2010) and population declines of Mauritian plants and animals (Jones et al. 1995; Safford 1997; Florens et al. 2017b). Today, < 2% of native forest cover remains and most of its endemic frugivores have disappeared, including the dodo, Raphus cucullatus. The island currently is left with only six endemic dispersers (Cheke and Hume 2008), of which only one can provide dispersal services for large-seeded, structurally important canopy trees (Florens et al. 2017a): the Mauritian flying fox (Pteropus niger Kerr, 1792). Pteropus niger feeds on the fruits of many native canopy tree species (Nyhagen et al. 2005; Florens et al. 2017a) that provide important services and resources to other plants and animals (Motala et al. 2007). The native forests therefore largely rely on the long-distance flight (Oleksy et al. 2019) and seed-dispersal abilities of P. niger for successful regeneration (Nyhagen 2004; Nyhagen et al. 2005; Florens et al. 2017a; Vincenot et al. 2017). The large number of IAS and lack of other large-bodied native seed dispersers make Mauritius an ideal platform to study the impact of IAS on flying foxes and their ecological roles. The invasive long-tailed macaque (Macaca fascicularis Raffles, 1821) was introduced in Mauritius ~400 years ago (Cheke and Hume 2008) and may threaten P. niger by competing for native fruits. Introduced, nonhominid primates have both directly and indirectly caused numerous species declines and extinctions on islands globally (Carter and Bright 2002; Kemp and Burnett 2003; Jones et al. 2018; Hanson et al. 2019). Although food competition between an introduced mammal and a native species only has been demonstrated for invasive rodents both on islands (Harris and Macdonald 2007) and the mainland (Mazzamuto et al. 2017), multiple studies suggest introduced monkeys may limit food to native species (Oliveira and Grelle 2012; Camarotti et al. 2015). In Mauritius, M. fascicularis destroys (Baider and Florens 2006; Florens 2015; Krivek 2017) and exploits unripe fruit (Sussman et al. 2011), potentially reducing the availability of native fruits to bats (Sussman et al. 2011; Florens 2015) and thereby decreasing seed dispersal. Exclusion experiments often are needed to demonstrate interspecific competition but are usually limited to studies on small animals (Petren and Case 1996; Cadi and Joly 2003; Duyck et al. 2006; Jensen et al. 2006). For study systems such as ours, a necessary first step is to quantify dietary overlap between species and explore the potential competitive advantage of one species over the other for a shared limited food source (Stier and Mildenstein 2005). Forest fruits may be a limited resource because native forests are scarce, and M. fascicularis may have a competitive advantage over P. niger by being able to exploit unripe fruits. Potential competition for native fruits with M. fascicularis and lack of native habitat may have led P. niger to feed on commercially valuable orchard crops in plantations and private backyards, where it is now considered a pest and subjected to large-scale culling (Vincenot et al. 2017; Oleksy et al. 2021; Tollington et al. 2019). Pteropus spp. shift to a diet mainly consisting of cultivated fruits when native forest resources are low (Parry-Jones and Augee 1991; Banack 1998), thereby likely allowing them to survive in fragmented landscapes (Stier and Mildenstein 2005; Jenkins et al. 2007; Luskin 2010). However, they still can depend greatly on native forest fruits (Stier and Mildenstein 2005; Javid et al. 2017), possibly because of the higher nutritional value of native compared to commercial fruits (Nelson et al. 2000). In addition, some Pteropus spp. prefer native over cultivated fruits (Parry-Jones and Augee 1991; Banack 1998). Therefore, if M. fascicularis reduces the availability of scarce, nutritionally important and preferred native food sources for P. niger, it will likely fuel the persecution of bats due to increased damage to commercial fruit. Persecution has made conservation of P. niger extremely challenging. Exaggerated and sometimes false claims about bats’ damage to commercial fruits, and the subsequent pressure on the government to protect fruit growers’ businesses, have resulted in a lack of evidence-based decision-making (Florens and Vincenot 2018; Florens and Baider 2019). For example, the cull in 2018 was carried out despite evidence for the ineffectiveness of culling in boosting fruit growers’ profits, while ignoring the increased extinction risk to the endemic P. niger (Florens and Vincenot 2018; Florens and Baider 2019). The culls are unlikely to stop in the near future, illustrated by the fact that the government announced another cull in October 2020 (L’Express 2020). Therefore, it is not only increasingly important to educate consumers, fruit growers, and policymakers (Florens and Baider 2019), but also to start addressing the potential role of M. fascicularis in this conflict. Here, we aimed to understand dietary overlap between M. fascicularis and P. niger when feeding on native fruit, and the implications of this overlap. Hence, we (a) investigated the diet of both species; (b) determined the influence of tree characteristics on fruit consumption by P. niger and M. fascicularis; (c) empirically quantified how M. fascicularis affects the abundance of native fruits; and (d) inferred possible impacts on seed dispersal by P. niger and consequent forest regeneration. We hypothesized that M. fascicularis reduce the abundance of native fruits eaten by P. niger due to dietary overlap. Because tall trees are more accessible to flying foxes and likely harder to climb for macaques than small trees, we also predicted that P. niger eat a larger proportion of fruits on tall trees, whereas M. fascicularis eat a larger proportion of fruits on small trees. We then discuss how our findings could improve conservation of P. niger and enhance its functional role in the native remnant forests. Materials and Methods Study area The study was carried out in three wet forest and two dry forest sites in Mauritius (Fig. 1). The sites are located within two of three major forest areas on the island (Moka Range and the Black River Gorges National Park) that are inhabited by M. fascicularis (Sussman et al. 2011) and used by P. niger for roosting and feeding (Oleksy et al. 2019). The sites were selected for their abundance of large endemic and native canopy trees and are similar in species composition to other native remnant forests, and thus are representative of potential conflict zones between P. niger and M. fascicularis. Two of the three wet forest sites were in Brise Fer in the Black River Gorges National Park (BRGNP): the closed management area, Brise Fer CMA (−20.376°N, 57.443°W), and a patch of weeded forest southeast of the CMA, here referred to as “Brise Fer semi-restored” (−20.385°N, 57.452°W). Both Brise Fer sites are at 600 m a.s.l. and receive < 4,500 mm of rainfall per year (Sevathian and Atkinson 2007). The forest in Brise Fer CMA was weeded in the late 1980s and exotic plants since have been removed frequently to prevent them from dominating the understory. These actions have resulted in a tall native canopy including some of the oldest trees in Mauritius today. Weeding started in the forest in Brise Fer semi-restored 10 years after the start of weeding in Brise Fer CMA. As a result, the canopy is more open in Brise Fer CMA. The final wet forest site was in a remnant wet forest on Calebasses Mountain (−20.181°N, 57.585°W), a mountainous area in the Moka Range (500–600 m a.s.l.) with rainfall of 2,500–4,500 mm/year. The dry forest sites were in the Lower Gorges of the BRGNP (−20.389°N, 57.433°W) and in Bon Amour (−20.145°N, 57.583°W). Both dry forest sites are 150–300 m a.s.l. and receive < 1,500 mm of rainfall/year. Fig. 1. Open in new tabDownload slide Mauritius on the east of Madagascar (top left) and closer view of the five study sites in Mauritius. Fig. 1. Open in new tabDownload slide Mauritius on the east of Madagascar (top left) and closer view of the five study sites in Mauritius. Dietary overlap between P. niger and M. fascicularis To assess the impact of M. fascicularis on food also eaten by P. niger, and on reproduction of native plant species, we measured the fruit drop under the canopy of 17 canopy tree species known to be included in P. niger’s diet (Florens et al. 2017a: appendix 1) from February 2018 until January 2019. We used seed traps to sample fruits and flowers that dropped from the canopy. The seed traps were constructed by stretching a 1 × 1 m quadrat of water-permeable mesh between plant stems with a string at all four corners. The quadrat was raised 30 cm above the ground to minimize fruit removal by rats. Three seed traps were placed under three to six individuals of the 17 focal species at each study site (Supplementary Data SD1). We recorded diameter at breast height (DBH) for every replicate as an index of tree size. To determine the start of flowering, focal trees were thoroughly searched for flowers on a monthly basis either by using binoculars or by climbing with climbing harness and tapes. Phenophases such as flowering and fruiting can be asynchronous (Borchert 1983) because rainfall is an important cue for fruiting in tropical trees (Mendoza et al. 2017) and varies greatly across sites. Start of flowering or fruiting of species occurring at multiple sites therefore was confirmed at every site. We then checked seed traps on a weekly basis from the start of the flowering until the end of the fruiting period for each tree. All fruits and flowers (intact and eaten), seeds (intact and eaten), bat ejecta pellets (compressed fibrous material that remains after P. niger squeezes the juice from the flesh; Supplementary Data SD2), and feces of various species, were recorded for the seed traps. Fruits eaten by P. niger were identified by their typical triangular-shaped canine imprint (Banack 1998; McConkey and Drake 2015), contrary to the larger rounded canine imprints and large incisor marks made by M. fascicularis (Krivek 2017; Supplementary Data SD2). Consumption of fruits by invasive rats could be recognized by the gnaw marks left on fruits by the distinctive upper and lower incisors. Fruits eaten by parrots (either native Psittacula eques or invasive Psittacula krameri) could be distinguished by triangular marks left by the under and upper mandibles (VSG 2018). One limitation of this method was that ingested fruits, intact fruits dropped by animals, or fruit stored in cheek pouches could not be detected, which, for example, may occur when M. fascicularis handles the fruits of small-seeded tree species selected for this study (Cossinia pinnata and Syzygium glomeratum) that could easily be ingested or stored in its cheek pouches. We also examined each fruit sampled by seed traps for seed maturity by opening the fruit and extracting the seed. Mature seeds were hard and fully developed, whereas immature seeds were soft and empty inside. Ripe fruits were characterized by soft flesh and a sweet smell, while unripe or premature fruits were typically tough and had soft seeds that could easily be squashed between thumb and index finger. Both intact fruit and fruit eaten by M. fascicularis and P. niger were totaled for every focal species to determine the percentage of fruit that was eaten by both animals. To confirm the identity of the animals that were visiting the studied trees, a single camera trap was attached to the trunk of one tree replicate for every species and aimed at the canopy or feeding branches. Five camera traps were used, consisting of a combination of H3 8 MP (Shenzhen VERZON E-Business Co. Ltd., Shenzhen, China) and Akaso 12 MP (Akaso, Frederick, Maryland) wildlife cameras. A single 10-s video was recorded whenever motion was detected. Camera traps often would remain functional only for 1 to 3 days, and footage of animals feeding on fruits could only be collected for Cassine orientalis, Diospyros leucomelas, and Foetidia mauritiana. The footage provided evidence for linking various bite mark types to species and insights into fruit handling by P. niger, M. fascicularis, and R. rattus (Supplementary Data SD3). Characteristics of feeding trees To determine features of trees where P. niger and M. fascicularis were the most common frugivores, the effect of DBH on the proportion of fruits eaten by each species was analyzed with logistic regression models in R (R Core Team 2017). We also explored relationships between the response variables and other potential predictors, and included those with R ≥ 0.1. As a result, the following explanatory variables were selected for both models: species, site, and DBH. Based on the second-order Akaike Information Criterion (AICc), the most parsimonious model was selected out of several models including all possible explanatory variable combinations. For the most parsimonious model of feeding tree use by P. niger and M. fascicularis (Table 1), the response variable caused quasi-complete separation for the site and species predictors, leading to inflated coefficients. We therefore used penalized logistic regression to produce models with more reasonable coefficients (bias reduction of maximum likelihood estimates using the function brglm from package brglm—Kosmidis 2019). Post hoc contrast tests were carried out for the pairwise comparisons between sites, adjusting P-values for multiple comparisons using the Tukey method (function emmeans from package emmeans—Lenth 2019). Table 1. AIC results for a priori models explaining tree selection by Pteropus niger and Macaca fascicularis. The proportion of fruits eaten on trees by P. niger (proportion bat) and M. fascicularis (proportion macaque) were response variables, and diameter at breast height (DBH) of trees, site, and species were used as predictors. Selection of the most parsimonious model was based on the second-order Akaike Information Criterion (AICc). The number of parameters (k), difference between model AICc and lowest AICc in the model set (∆AICc), and AIC weight (ω), also are provided for every model. . k . ∆AICc . ω . Model: tree selection by Pteropus niger  Proportion bat ~ Species + Site 17 0.0 0.8  Proportion bat ~ DBH + Species + Site 18 2.6 0.2  Proportion bat ~ DBH + Species 14 151.5 9.9 * 10−34  Proportion bat ~ DBH + Site 6 3,967.4 0.0  Null model 1 4,832.4 0.0 Model: tree selection by Macaca fascicularis  Proportion macaque ~ DBH + Species + Site 18 0.0 0.8  Proportion macaque ~ DBH + Species 14 3.6 0.1  Proportion macaque ~ Species + Site 17 7.3 0.02  Proportion macaque ~ DBH + Site 6 2,775.5 0.0  Null model 1 3,457.2 0.0 . k . ∆AICc . ω . Model: tree selection by Pteropus niger  Proportion bat ~ Species + Site 17 0.0 0.8  Proportion bat ~ DBH + Species + Site 18 2.6 0.2  Proportion bat ~ DBH + Species 14 151.5 9.9 * 10−34  Proportion bat ~ DBH + Site 6 3,967.4 0.0  Null model 1 4,832.4 0.0 Model: tree selection by Macaca fascicularis  Proportion macaque ~ DBH + Species + Site 18 0.0 0.8  Proportion macaque ~ DBH + Species 14 3.6 0.1  Proportion macaque ~ Species + Site 17 7.3 0.02  Proportion macaque ~ DBH + Site 6 2,775.5 0.0  Null model 1 3,457.2 0.0 Open in new tab Table 1. AIC results for a priori models explaining tree selection by Pteropus niger and Macaca fascicularis. The proportion of fruits eaten on trees by P. niger (proportion bat) and M. fascicularis (proportion macaque) were response variables, and diameter at breast height (DBH) of trees, site, and species were used as predictors. Selection of the most parsimonious model was based on the second-order Akaike Information Criterion (AICc). The number of parameters (k), difference between model AICc and lowest AICc in the model set (∆AICc), and AIC weight (ω), also are provided for every model. . k . ∆AICc . ω . Model: tree selection by Pteropus niger  Proportion bat ~ Species + Site 17 0.0 0.8  Proportion bat ~ DBH + Species + Site 18 2.6 0.2  Proportion bat ~ DBH + Species 14 151.5 9.9 * 10−34  Proportion bat ~ DBH + Site 6 3,967.4 0.0  Null model 1 4,832.4 0.0 Model: tree selection by Macaca fascicularis  Proportion macaque ~ DBH + Species + Site 18 0.0 0.8  Proportion macaque ~ DBH + Species 14 3.6 0.1  Proportion macaque ~ Species + Site 17 7.3 0.02  Proportion macaque ~ DBH + Site 6 2,775.5 0.0  Null model 1 3,457.2 0.0 . k . ∆AICc . ω . Model: tree selection by Pteropus niger  Proportion bat ~ Species + Site 17 0.0 0.8  Proportion bat ~ DBH + Species + Site 18 2.6 0.2  Proportion bat ~ DBH + Species 14 151.5 9.9 * 10−34  Proportion bat ~ DBH + Site 6 3,967.4 0.0  Null model 1 4,832.4 0.0 Model: tree selection by Macaca fascicularis  Proportion macaque ~ DBH + Species + Site 18 0.0 0.8  Proportion macaque ~ DBH + Species 14 3.6 0.1  Proportion macaque ~ Species + Site 17 7.3 0.02  Proportion macaque ~ DBH + Site 6 2,775.5 0.0  Null model 1 3,457.2 0.0 Open in new tab Seasonal variation in diet and consumption of plant parts and unripe fruit To determine other feeding sources used by P. niger and M. fascicularis and the type of plant parts consumed, a 4 by 500 m line transect was established at every site. We walked the transects slowly (1 km/h) every month from May 2018 until January 2019. The ground was actively searched and scanned for eaten plant parts, and the following parameters were recorded: food plant species, type of plant part consumed (leaves, flowers, unripe, and ripe fruit), and type of animal that had consumed the plant part (flying fox, macaque, rat, or parrot). We then calculated the proportions of native versus exotic food species and ripe versus unripe fruit eaten by P. niger and M. fascicularis. In addition, the food species overlap between P. niger and M. fascicularis was calculated with pairwise Jaccard’s coefficients (Krebs 1989). A limitation of this method was that plant items completely swallowed by an animal could not be detected. Results Feeding patterns and fruit production Only the fruits of 13 of the 17 selected tree species were consumed by P. niger, M. fascicularis, or other species (Fig. 2). We sampled 13,785 fruits for these species across 102 trees in five sites (median [interquartile range (IQR)]: 11.5 [0.3–89.0] fruits per tree replicate). Of the sampled fruits, 2,038 (15%, 0 [0–4.8]) were eaten by flying foxes, 532 (4%, 0 [0–2.8]) by macaques, 5,436 remained intact (39%, 3.0 [0–19.8]), and 5,801 were eaten or parasitized by other animals (42%, 0 [0–3.0]). Cassine orientalis (n = 3,274, 179.0 [2.8–431.0]), S. glomeratum (n = 3,100, 133.0 [40.0–311.5]), and Terminalia bentzoë (n = 4,992, 1,210.0 [1,006.0–2,095.0]) produced the largest numbers of fruit and collectively accounted for 83% of all recorded fruit. Furthermore, C. orientalis (n = 2,044, 71.0 [1.3–225.0]) and S. glomeratum (n = 2,037, 97.5 [14.5–180.8]) accounted for 75% of all recorded dropped intact fruits, and T. bentzoë accounted for 75% of fruits eaten or parasitized by other animals (n = 4,365, 1,089.0 [720.5–2,006.5]). Terminalia bentzoë was exclusively parasitized by moth larvae. Rattus rattus mainly accounted for the remaining 25% of fruits eaten or parasitized by other animals across other species (n = 1,134, 19.0 [3.8–42.8]), and mostly consumed C. orientalis (n = 1,096, 40.5 [0.5–69.3]). Macaca fascicularis consumed variable portions of fruit for seven out of 13 species, including 90–95% of fruits of Diospyros tesselaria (n = 364, 0 [0–18.0]), 85–100% of D. leucomelas (n = 44, 9.0 [7.0–12.0]), 20–21% of Labourdonnaisia glauca (n = 37, 2.5 [0–9.3]), 23–96% of Labourdonnaisia revoluta (n = 24, 0 [0–3.0]), 28% of Mimusops maxima (n = 26, 3.0 [2.0–6.0]), 20% of Mimusops petiolaris (n = 18, 1.5 [0.8–0.3]), and 50% of Sideroxylon grandiflorum (n = 7, 1.0 [0–2.0]) across the sites (Fig. 2). Fig. 2. Open in new tabDownload slide Percentage of sampled fruits that remained intact or were eaten by either Pteropus niger, Macaca fascicularis or other animals (including parasites) for Diospyros leucomelas (Dio leu), Foetidia mauritiana (Foe mau), Mimusops petiolaris (Mim pet), Labourdonnaisia glauca (Lab glau), Mimusops maxima (Mim max), Labourdonnaisia revoluta (Lab rev), Stadmania oppositifolia (Sta opp), Sideroxylon grandiflorum (Sid gra), Syzygium glomeratum (Syz glo), and Terminalia bentzoë (Ter ben) in Bon Amour (A), Brise Fer CMA (B), Brise Fer semi-restored (C), Lower Gorges (D), and Calebasses Mountain (E). Species for which no eaten fruits were recorded have been omitted. Fig. 2. Open in new tabDownload slide Percentage of sampled fruits that remained intact or were eaten by either Pteropus niger, Macaca fascicularis or other animals (including parasites) for Diospyros leucomelas (Dio leu), Foetidia mauritiana (Foe mau), Mimusops petiolaris (Mim pet), Labourdonnaisia glauca (Lab glau), Mimusops maxima (Mim max), Labourdonnaisia revoluta (Lab rev), Stadmania oppositifolia (Sta opp), Sideroxylon grandiflorum (Sid gra), Syzygium glomeratum (Syz glo), and Terminalia bentzoë (Ter ben) in Bon Amour (A), Brise Fer CMA (B), Brise Fer semi-restored (C), Lower Gorges (D), and Calebasses Mountain (E). Species for which no eaten fruits were recorded have been omitted. Footage from camera traps indicated that P. niger was the most frequent forager on F. mauritiana (110 of 147 [75%] videos from 28 May 2018 to 7 June 2018). All F. mauritiana fruits eaten by P. niger that were recorded in the seed traps showed triangular canine marks, were cleared of most fiber, and the seeds remained undamaged. Ripe fruits eaten by P. niger and recorded in the seed traps for L. glauca, M. maxima, Stadmania oppositifolia, and S. glomeratum also typically were cleared of fruit pulp and seeds remained undamaged. Conversely, footage indicated M. fascicularis was the only feeder on D. leucomelas (all six videos on 15 April 2018), and M. fascicularis takes a single bite of the unripe fruit, often dropping a large chunk of the fruit with underdeveloped and damaged seeds. Unripe fruits eaten by M. fascicularis recorded in the seed traps for D. leucomelas, M. maxima, M. petiolaris, L. glauca, L. revoluta, D. tesselaria, and S. grandiflorum, had a single bite mark, typically resulting in destruction of premature seeds. As a result, at least half of pulp was not consumed. In addition, camera trap footage indicated that R. rattus was the main feeder on C. orientalis (44 of 44 videos from 30 October 2018 to 13 November 2018). Fruiting patterns and fruit availability The start of fruiting for C. orientalis, D. tesselaria, L. revoluta, Protium obtusifolium, S. glomeratum, and S. oppositifolia, varied among study sites (Supplementary Data SD4). For seven of the nine species P. niger visited during the study, P. niger only fed on fruits when ripe fruit was available (Supplementary Data SD4). The other two species were fed on when only unripe fruit was available. For seven of nine species, P. niger did not feed on fruits during all months that ripe fruit was available (Supplementary Data SD4). For example, P. niger fed on C. orientalis in Brise Fer CMA and T. bentzoë in Bon Amour only during 1 and 2 months, respectively, even though ripe fruits were available for 4 and 11 months, respectively (Supplementary Data SD4). Furthermore, fruits of C. orientalis and S. oppositifolia only were consumed by P. niger in a single site, even though these species were present in other sites (Supplementary Data SD4). In contrast, M. fascicularis started feeding before ripe fruit became available for six out of seven species visited during our study. Characteristics of feeding trees The top-ranked model for explaining relative fruit consumption by P. niger included tree species and site but omitted DBH (Table 1). There was extensive variation in the proportion of fruits consumed across sites (Supplementary Data SD5). Post hoc analysis revealed that the proportion of fruits eaten by P. niger was higher in Brise Fer CMA compared to Calebasses Mountain and Brise Fer semi-restored (P < 0.05; Supplementary Data SD5), suggesting the degree of forest invasion could play a role in feeding tree selection by P. niger. However, the model fit was poor (807.68 residual deviance with 53 d.f.), indicating overdispersion. The model was run again with a quasi-binomial distribution to account for overdispersion, which resulted in smaller coefficient estimates but no substantial change to our P-values. In summary, our model shows P. niger eats a larger proportion of fruits on trees in fully restored compared to some degraded forests. The top-ranked model for explaining relative fruit consumption by M. fascicularis included tree species, site, and DBH (Table 1). The proportion of fruits consumed by M. fascicularis decreased for trees with larger DBH (Supplementary Data SD6). The model fit was good (95.64 residual deviance with 52 d.f.). Seasonal variation in diet and consumption of plant parts and unripe fruit Additional plant species eaten by P. niger and M. fascicularis were identified during the monthly transect walks (Supplementary Data SD7 and SD8). Pteropus niger mostly ate ripe fruits, while M. fascicularis mainly consumed unripe fruits or seeds (Table 2). Furthermore, P. niger exclusively consumed leaves of four out of the 23 identified food species and exclusively consumed flowers of D. tesselaria (Table 2). There was high overlap in food species used throughout the year by P. niger and M. fascicularis (Table 2). Two shared food species were used during completely different periods by P. niger and M. fascicularis. For M. petiolaris, P. niger ate fruits during May–July, whereas M. fascicularis used this food species during March–May (Supplementary Data SD7 and SD8). For L. revoluta, P. niger ate the fruits during August, whereas M. fascicularis used this food species during July and October (Supplementary Data SD7 and SD8). Table 2. Number of food species eaten by Pteropus niger and Macaca fascicularis and overlap of food species between both mammal species. Included are the number of exotic and native-endemic species both animals fed on and the number of species for which they exclusively consumed ripe fruit, unripe fruit, flowers, or leaves, and the number of species exclusively consumed by either animal. Food species overlap was calculated with a pairwise Jaccard’s coefficient: J = A/(A + B + C), where A is the total number of species eaten by both P. niger and M. fascicularis, B the species only eaten by P. niger and C species only eaten by M. fascicularis. No. of food species . Pteropus niger . Macaca fascicularis . Total 23 16 Native-endemic 19 12 Exotic 4 4 Exclusively ripe fruit eaten 12 4 Exclusively unripe fruit eaten 3 8 Exclusively flowers eaten 1 1 Exclusively leaves eaten 4 0 Exclusively eaten by either animal 12 5 Food species overlap 62.2% No. of food species . Pteropus niger . Macaca fascicularis . Total 23 16 Native-endemic 19 12 Exotic 4 4 Exclusively ripe fruit eaten 12 4 Exclusively unripe fruit eaten 3 8 Exclusively flowers eaten 1 1 Exclusively leaves eaten 4 0 Exclusively eaten by either animal 12 5 Food species overlap 62.2% Open in new tab Table 2. Number of food species eaten by Pteropus niger and Macaca fascicularis and overlap of food species between both mammal species. Included are the number of exotic and native-endemic species both animals fed on and the number of species for which they exclusively consumed ripe fruit, unripe fruit, flowers, or leaves, and the number of species exclusively consumed by either animal. Food species overlap was calculated with a pairwise Jaccard’s coefficient: J = A/(A + B + C), where A is the total number of species eaten by both P. niger and M. fascicularis, B the species only eaten by P. niger and C species only eaten by M. fascicularis. No. of food species . Pteropus niger . Macaca fascicularis . Total 23 16 Native-endemic 19 12 Exotic 4 4 Exclusively ripe fruit eaten 12 4 Exclusively unripe fruit eaten 3 8 Exclusively flowers eaten 1 1 Exclusively leaves eaten 4 0 Exclusively eaten by either animal 12 5 Food species overlap 62.2% No. of food species . Pteropus niger . Macaca fascicularis . Total 23 16 Native-endemic 19 12 Exotic 4 4 Exclusively ripe fruit eaten 12 4 Exclusively unripe fruit eaten 3 8 Exclusively flowers eaten 1 1 Exclusively leaves eaten 4 0 Exclusively eaten by either animal 12 5 Food species overlap 62.2% Open in new tab Discussion Dietary overlap between P. niger and M. fascicularis Overall, fruit production across all tree replicates was low and highly variable as indicated by the low median relative to the high IQR (11.5 ± 88.8). Even though macaques only ate ~4% of the fruits recorded during this study, these fruits accounted for 20% to 100% of total fruit availability of half of the selected tree species visited by either P. niger or M. fascicularis. The total overlap in food species between P. niger and M. fascicularis across all sites was 62.2%, indicating that M. fascularis may reduce fruit availability for most food species used by P. niger. Moreover, macaques ate 90–95% of D. tesselaria fruits. This species represents one of the more common fruiting trees in native remnant forests and may be a more important resource on an island scale than most of the other species. Nevertheless, 39% of recorded fruits were intact, and we did not record macaques eating fruits for species that produced the most fruit (C. orientalis and S. glomeratum), suggesting fruit may not necessarily be limited for either mammal species. This could be partly explained by the limitations of our methods, because collateral damage (animals accidentally or intentionally dropping intact fruit) and ingestion of fruits by M. fascicularis, could not be recorded. Syzygium glomeratum and C. orientalis had the smallest fruits out of our studied species, meaning M. fascicularis may simply have ingested them whole. Furthermore, field observations revealed that a single macaque can deplete large quantities of fruit on a tree (invasive Litsea glutinosa) within a single feeding session by plucking and dropping intact fruit. This behavior first was described by Thompson (1880), who documented that M. fascicularis throws down unripe fruit from all structurally important forest trees, reducing the availability of viable seeds for eventual germination. Given the limitations of our camera traps, we failed to record this potential feeding behavior. Consequently, we can assume the actual fruit drop by M. fascicularis may be significantly higher, and its diet may be broader, than presented in this paper. Pteropus niger ate more ripe fruit than unripe fruit. This is common in Pteropus spp. (Bollen and Van Elsacker 2002; Luft et al. 2003) because they depend on their sense of smell to locate the food source (Hodgkison et al. 2003). The pulp of ripe fruits treated by P. niger usually was removed completely. Contrary to P. niger, M. fascicularis mainly consumed fruit at an unripe stage and often would fail to remove at least half of the (still very tough) fruit pulp. Pulp removal often is necessary to enhance germination of the seed (Izhaki and Safriel 1990; Barnea et al. 1991; Traveset 1998; Fedriani et al. 2012), suggesting that P. niger is a more effective disperser than M. fascicularis. The large number of unripe fruits damaged by macaques, and the lack of fruits eaten by P. niger for Diospyros boutoniana (Endangered), D. nodosa (Critically Endangered), D. leucomelas, D. tesselaria, L. revoluta, and M. petiolaris suggest M. fascicularis may deplete unripe fruits on trees before P. niger can use them. As a result, M. fascicularis may restrict P. niger’s diet, resulting in some degree of competition. Furthermore, this also might disrupt effective seed dispersal by P. niger including rare and threatened canopy species. In Macaca spp., consumption of unripe fruit and seeds often is considered a response to low fruit availability (Huang et al. 2015; Tang et al. 2016). In regions with sufficient fruit sources, Macaca spp. typically prefer ripe over unripe fruit and even can be effective seed dispersers (Yeager 1996; Sengupta et al. 2014; Sengupta and Radhakrishna 2015). Our results show that M. fascicularis feeds mainly on unripe fruits, possibly indicating low fruit availability. However, this scenario is contradicted by the fact that M. fascicularis ate a small proportion of fruit overall. Nevertheless, accurately quantifying fruit consumption by M. fascicularis may be possible only by directly observing individuals or using high-quality camera traps. Therefore, additional research is required to test the hypotheses that M. fascicularis limits the availability of native fruits to P. niger or that native fruits are limiting for either species. We suggest that long-term observational studies on M. fascicularis diet through habituation and tracking of macaque groups in native forest remnants paired with fruit availability assessments will be valuable. Furthermore, P. niger exclusively fed on leaves of certain species during most of the study, suggesting leaves might be an important dietary supplement. In vertebrates, strictly frugivorous diets are rare because of the low amount of protein in fruits (Izhaki and Safriel 1989). As a result, some frugivorous mammals, including flying foxes, supplement their diets with protein-rich leaves (Jordano 2000; Nelson et al. 2005; Ganzhorn et al. 2009). It is unlikely that feeding on leaves by P. niger is a result of food scarcity, because ripe fruits of certain species (e.g., C. orientalis and T. bentzoë) were available during these months and at times were completely ignored by P. niger. Nutrient studies would be necessary to reveal why P. niger uses certain food sources (leaves, flowers, fruit) and ignores others. However, for C. orientalis, R. rattus may have prevented P. niger from visiting the trees because R. rattus ate most C. orientalis fruits in all sites and can be a nocturnal feeder like P. niger. Therefore, R. rattus also may reduce the availability of certain native fruits and influence food choice by P. niger. Role of remnant forests in food availability Six of the 13 canopy species that P. niger used differed in fruiting periods among sites. It is unlikely that the start of flowering was missed in any of the sites because the trees were checked thoroughly for fruits and flowers on a monthly basis. Asynchronous fruiting patterns within Diospyros spp. also were observed between slopes facing different directions within the same site. Asynchronous flowering patterns can be driven by variation in timing of leaf fall, which can be determined by water availability and rainfall (Reich and Borchert 1984). As a result, the timing of flowering caused by leaf fall can be highly asynchronous within tropical tree species (Borchert 1983). Annual rainfall varies greatly across Mauritius and many tree species occupy both wet and dry areas with diverse microclimates, so asynchronous fruiting is expected to be common. A given site therefore may provide fruits of certain species during a time of the year when other sites do not. For example, C. orientalis started fruiting 3 months later in Bon Amour than in the Lower Gorges, meaning ripe fruits are likely to still be available in Bon Amour after C. orientalis has finished fruiting in the Lower Gorges. This may become increasingly important in the face of climate change because rainfall variability is expected to increase in the tropics (Sakai and Kitajima 2019). As a result, fruiting patterns could become more variable (Dunham et al. 2018) and remnant forests across the island may help maintain a steady supply of different fruits to P. niger throughout the year. Pteropus niger can travel far to feed on specific resources, potentially moving over the entire island and feeding in different locations within a single night (Oleksy et al. 2019). Characteristics of feeding trees Interestingly, DBH was omitted from the most parsimonious model; our hypothesis that P. niger eats a higher proportion of fruits on taller trees thus was rejected. Pteropus niger has excellent climbing abilities, allowing them to maneuver between tall and small trees. This is well illustrated by our frequent sightings of P. niger feeding in small strawberry guava (P. cattleyanum) in the forests. A nearby tall tree could provide an access point to reach low canopy vegetation. Therefore, proximity of tall trees around the feeding tree may be a better predictor for feeding tree selection by P. niger than trunk size of the feeding tree. Furthermore, the proportion of fruits eaten by P. niger was higher in Brise Fer CMA than in Brise Fer semi-restored. These sites are similar in species composition, but the forest in Brise Fer semi-restored is more open and degraded than Brise Fer CMA and may differ in terms of abundance of fruiting trees. The proximity of other fruit sources therefore may influence the feeding tree selectivity by P. niger. In addition, increased alien plant invasion reduces the fruit production of native angiosperms (Monty et al. 2013; Krivek et al. 2020) and therefore affects visitation rates by P. niger (Krivek et al. 2020). Finally, the most parsimonious model explaining relative fruit consumption by P. niger was overdispersed, indicating other unmodeled factors likely explain feeding tree selection. The proportion of fruits eaten by M. fascicularis was higher on trees with smaller DBH compared to trees with larger DBH, confirming our hypothesis that M. fascicularis eats a larger proportion of fruits on smaller trees. Large straight trunks are difficult for M. fascicularis to climb. To reach the canopy of a large tree the macaques therefore would have to use smaller trees as stepping stones. However, the accessibility to M. fascicularis is not only explained by DBH because DBH does not correlate always with tree height. Proximity of the canopies of surrounding trees and trunk shape also will affect accessibility. Nevertheless, our analysis shows that smaller trees may have a higher risk of having all their fruits depleted at an unripe stage by macaques than larger trees, meaning seed dispersal for smaller trees may be reduced. Impact of M. fascicularis on forest regeneration Although M. fascicularis may prevent the development of viable seeds of some plant species by destroying unripe fruit, successful regeneration of these species is not guaranteed, even if M. fascicularis is excluded from forests. Young plants will remain extremely vulnerable during the growth stages that follow the seed stage. The transition from seed to seedling and the subsequent crucial survival of seedlings likely is limited by exotic plant invasion (Lorence and Sussman 1986; Kitajima and Fenner 2000; Florens et al. 2017b). Except for the restored forest in Brise Fer CMA, our study sites had high levels of exotic plant invasion. Therefore, besides mitigation of fruit destruction by M. fascicularis, weed removal will be crucial to promote forest regeneration (Baider and Florens 2011; Florens et al. 2017b). If consumption of unripe fruit is a result of fruit scarcity, M. fascicularis may consume more ripe fruit and thus act as an effective seed disperser when forest resources are abundant. However, high birth rate, lack of natural predators, and the ability to exploit both unripe fruit and anthropogenic food sources (Bertram 1994; Sussman et al. 2011) suggest the population of M. fascicularis could exceed carrying capacity of the forest eventually. Implications for the conservation of P. niger and native forests Although exploitative competition typically is proven via exclusion experiments, which would be logistically difficult in this system, our results suggest that M. fascicularis potentially reduces the availability of native fruit of certain threatened tree species to P. niger and even halts regeneration of some tree species. However, more empirical research is required to demonstrate whether availability of native fruits to P. niger is limited by M. fascicularis. Nevertheless, our findings highlight the potential detrimental consequences of introduced frugivores on keystone seed dispersers and their habitat. These findings emphasize that an integrative approach to forest restoration, requiring both systematic weeding efforts to rid native remnant forests of invasive plants, and an understanding of seed predators and restoration of plant–disperser interactions, may be valuable to conservation of P. niger and its habitat. Our study also showed that food selection by flying foxes and the role of remnant forests in food provision during different seasons are research gaps that need attention to inform conservation efforts across Mauritius. Exploring these gaps may become increasingly urgent given climate change, because phenology patterns are likely to change, and native forests will continue to face high rates of degradation by invasive plants. Addressing these knowledge gaps also will be crucial to reducing persecution of bats by humans. Restoration of remnant forest fragments may increase availability of native fruits to P. niger, potentially minimizing the impact of P. niger on commercial orchards, and assisting with a long-term solution to the current persecution of bats at commercial fruit plantations in Mauritius. We therefore suggest appointing core areas for restoration in the Moka Range, because few to no restoration efforts are underway in this region even though it contains high-quality, native remnant forests that can provide suitable habitat to P. niger. Acknowledgments We thank the Rufford Small Grants Foundation (grant 23082-1) for their financial support. We also thank the National Parks and Conservation Service, the Forestry Service, and the Ministry of Agro Industry and Food Security for providing permits to work on state land and in the Black River Gorges National Park. We are particularly grateful to C. Baider of The Mauritius Herbarium and F. B. V. Florens of the University of Mauritius for their plant identification support and for sharing their enthusiasm for ecology and the Mauritian flora and fauna. We also thank I. Janoo and L. Ayady of the Ecosystem Restoration Alliance, Indian Ocean (ERA), for exceptional assistance with field activities. Moreover, we are grateful to M. Beaumont of the University of Bristol for assistance with data analyses. We thank two anonymous reviewers for their helpful comments. No potential conflict of interest was reported by the authors. Supplementary Data Supplementary data are available at Journal of Mammalogy online. Supplementary Data SD1.—All species selected for the competition assessment. The number of replicates per species is listed for all study sites: Bon Amour (BA), Brise Fer CMA (BFC), Brise Fer semi-restored (BFs), Lower Gorges (LG), and Calebasses Mountain (CM). The hyphen indicates species-absence in a site. Supplementary Data SD2.—Ejecta pellets and fruit of Mimusops maxima eaten by Pteropus niger showing triangular canine marks (left A) and fruit of M. maxima eaten by Macaca fascicularis showing a rounded incisor mark (right A). Food items consumed by P. niger, from left to right: Labourdonnaisia glauca, M. maxima, Hugonia serrata, Psidium cattleyanum, and Syzygium glomeratum (B). Unripe fruit and seed damaged by M. fascicularis: Diospyros tesselaria (top C) and Diospyros nodosa (bottom C). Macaca fascicularis incisor marks can be seen on D. tesselaria fruits (top C). Supplementary Data SD3.—Camera trap footage of Pteropus niger about to grab ripe Foetidia mauritiana fruit (left), Macaca fascicularis biting unripe Diospyros leucomelas fruit (middle) and Rattus rattus carrying unripe fruit of Cassine orientalis (right). Supplementary Data SD4.—The availability of ripe fruit for all 17 species selected for this study from February 2018 until January 2019 in all sites: Canarium paniculatum (Can pan), Cassine orientalis (Cas ori), Diospyros leucomelas (Dio leu), Diospyros tesselaria (Dio tes), Foetidia mauritiana (Foe mau), Labourdonnaisia glauca (Lab glau), Labourdonnaisia revoluta (Lab rev), Mimusops maxima (Mim max), Mimusops petiolaris (Mim pet), Protium obtusifolium (Pro obt), Sideroxylon puberulum (Sid pub), Sideroxylon grandiflorum (Sid gra), Stadmania oppositifolia (Sta opp), Syzygium duponti (Syz dup), Syzygium glomeratum (Syz glo), and Terminalia bentzoë (Ter ben), in Bon Amour (BA), Brise Fer CMA (BFC), Brise Fer semi-restored (BFs), Lower Gorges (LG), and Calebasses Mountain (CM). The gray horizontal bar represents the fruiting period during which only unripe fruits were available (light gray) and both unripe and ripe fruits were available (dark gray). The closed dots indicate during which months Pteropus niger consumed fruits and the open dots indicate during which months Macaca fascicularis consumed fruits. Supplementary Data SD5.—Regression coefficients, standard errors (SEs), and P-values (*P < 0.05, **P < 0.01, ***P < 0.001, NS = not significant), for the likelihood ratio tests (lr) for all predictors included in the logistic regression model best explaining tree selection by Pteropus niger. In addition, coefficients, SEs, and adjusted P-values for pairwise comparisons between sites, are provided (coefficients and SEs are given in log odds). Supplementary Data SD6.—Regression coefficients, standard errors (SEs), and P-values (*P < 0.05, **P < 0.01, ***P < 0.001, NS = not significant), for the likelihood ratio tests (lr) for all predictors included in the logistic regression model best explaining tree selection by Macaca fascicularis. In addition, coefficients, SEs, and adjusted P-values for pairwise comparisons between sites, are provided (coefficients and SEs are given in log odds). Supplementary Data SD7.—Species documented to have been eaten by Pteropus niger and the consumed plant parts (fruit = Fr, unripe = u, ripe = r, flower = Fl, leaves = Lv, and seeds = Se), across the five study sites: Bon Amour (BA), Brise Fer semi-restored (BFs), Brise Fer CMA (BFC), Lower Gorges (LG), and Calebasses Mountain (CM). Supplementary Data SD8.—Species documented to have been eaten by different animals (Type: Macaca fascicularis = M, Rattus rattus = R, Psittacula eques or Psittacula krameri = P) and the consumed plant parts (Eaten plant part: fruit = Fr, unripe = u, ripe = r, flower = Fl, and leaves = Lv) across the five study sites: Bon Amour (BA), Brise Fer CMA (BFC), Brise Fer semi-restored (BFs), Lower Gorges (LG), and Calebasses Mountain (CM). Literature Cited Aziz , S. A. , et al. 2017 . 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TI - First come, first served: fruit availability to keystone bat species is potentially reduced by invasive macaques
JF - Journal of Mammalogy
DO - 10.1093/jmammal/gyaa182
DA - 2021-02-15
UR - https://www.deepdyve.com/lp/oxford-university-press/first-come-first-served-fruit-availability-to-keystone-bat-species-is-Ds6XnIV1ff
SP - 1
EP - 1
VL - Advance Article
IS - 
DP - DeepDyve
ER -