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Pteropus rufus Food selection: exploited versus available food items In order to gain some insight into the flying foxes’ food selection, the different variables of several fruit and seed parameters were compared for the 175 available and 40 exploited food species (Table 4). Focusing on exploited food species only, it was apparent that mainly large trees and to a lesser extent small trees and shrubs are exploited for their fruits. No herbs and vines occur in the diet list. Berries are the fruit type most represented in the diet followed by drupes. Fruits with a strong odour are predominant in the diet, while all colours are present in approximately similar percentages. Fruits with many tiny seeds as well as one- to two-seeded fruits are well represented. In general fruit skin thickness is minimal and most fruits have a water content over 60%. Furthermore juicy fruits with a length between 10–30mm are most common in the list, but no particular fruit mass was most abundant. Seed length is often smaller than 10mm and seed mass less than 0.1g. The threshold for seed swallowing at our study site is as much as 10mm, with 4.4mm being the median diameter (N=38). Much of the differential use of fruits can be explained by a differential availability. There is only a significant difference between observed and expected values for the parameters pulp type, seed length and seed mass (Table 4). Fruits with juicy pulp are clearly preferred. Fibrous fruits and fruits without pulp, even though available, are not consumed by the flying foxes at all. Fruits with seed length smaller than 10mm are preferred to longer seeded-fruits. The most preferred seed mass is under 0.1g, but the 0.1–1.0g category still makes up one third of their food choice, while seeds heavier than 1g seem to be avoided. After sequential Bonferroni adjustment (Rice 1989 but see Moran 2003) none of these preferences remained significant. Germination trials Our faecal analyses show that seeds of at least 38 plant species pass through the digestive tract. Due to the scarcity of simultaneous presence of defecated seeds, control seeds and fruits, it was not always possible to obtain the same number of duplicates or the same number of seeds for all treatments. None of the defecated seeds looked damaged. Only five species provided enough seeds and fruits at the same time to start a germination experiment (Table 5). Passage through the digestive tract had neither a negative nor a positive impact on the germination rate and percentage of seeds germinated. It appears that seeds from intact control fruits take more time to germinate than seeds of faecal samples and control seeds. Numbers were too small, however, to allow statistical comparison. DISCUSSION Fruit diet Quantitatively The diet of P. rufus studied at Sainte Luce consists of 40 endemic plant species of the littoral forest. Even though our data set represents the most complete information available on the fruit diet of P. rufus in littoral forests today, it is probably an underestimation of their overall fruit diet for several reasons. First, by focusing mainly on faecal sample content, larger seeds, that are often spat out and less commonly eaten food species can be missed. Secondly, exotic species that are neither important nor typical for the littoral forest were omitted from our study. It is likely that the five seed species that could not be identified may represent seeds from such exotic species. They may also be fruit species eaten in other forest types and were as such not present in our 95
Chapter 3a reference collection. It cannot be excluded that the mountain rain forest, with a different floral composition (Koechlin et al. 1974) lies within the foraging range of the colony studied. Finally, there might also be an important temporal bias since a number of tree species in these littoral forests do not fruit annually but bi-annually or even less often (Randrihasipara pers. comm.; Bollen and Donati, Chapter 1). One year of sampling is not enough to establish the complete fruit diet of P. rufus. Nevertheless, our species accumulation curve indicates that a large proportion of the diet is indeed already known. Long-term studies are needed to further complete the diet list. Table 5. The number of weeks needed for the first germination (a) and the percentage of germinated seeds (b) for the three treatments: faecal seeds, control seeds, and control fruits. The values given here are median, average deviation (AD), and sample size (N). a) Germination rate Species Faecal seeds Control seeds Control fruits Median AD N Median AD N Median AD N Ludia antanosarum 2.5 - 1 3.5 0.5 2 7 - 1 Terminalia fatraea 25 - 1 22 0.9 3 26 1.2 5 Syzygium sp. 7 - 1 6 0.9 3 10 6.8 3 Ficus guatteriifolia 3.5 2.9 8 7 - 1 15.5 2.5 2 Rothmannia mandenensis 8 - 1 23 - 1 2 - 0 b) Percentage germinated Species Faecal seeds Control seeds Control fruits Median AD N Median AD N Median AD N Ludia antanosarum 25 - 1 62.5 12.5 2 33 - 1 Terminalia fatraea 60 - 1 30 15.6 3 30 8.0 5 Syzygium sp. 50 - 1 80 11.1 3 60 13.3 3 Ficus guatteriifolia 16 9 8 1 - 1 1 - 2 Rothmannia mandenensis 50 - 1 40 - 1 2 - 0 1 as these seeds are very tiny and numerous in fruits it is impossible to know exactly which percentage had germinated in both control seeds and control fruits. 2 ripe fruits fall apart in small pieces and therefore we could not sow complete fruits for comparison. Quantitative data on the diet of P. rufus in other parts of Madagascar are limited. In the gallery forest of Berenty the diet of P. rufus contains only 13 plant species, both flowers and fruits included (Long and Racey submitted). This much lower number can probably be best explained by the lower plant diversity of the much drier gallery forest. Racey et al. (in prep) have studied P. rufus in Madagascar for several years at different sites in Madagascar and their data on feeding ecology resulted in a diet list of 38 plant species for P. rufus with two thirds of these being fruit resources. Racey and Nicoll (1984) mention a fruit diet of 18 species for Pteropus seychellensis in the Seychelles and Parry- Jones and Augee (1991) found 22 fruit species in the diet of Pteropus poliocephalus. An extensive literature survey by Marshall (1983) resulted in a list of 62 plant genera consumed for their fruits by all Pteropus spp. (N=67) together. All these numbers demonstrate that our diet list including 40 species belonging to 28 genera is quite extensive. As for the quantitative data of our observations, we believe there may be a bias on the visitation length scored, as flying foxes would fly away when they detected our 96
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Fruit-frugivore interactions in a M
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Cover photographs: Front: Back: -
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Table of contents III. General conc
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Acknowledgements A special word of
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PREFACE Preface This work brings to
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General introduction Finally, seed
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General introduction high as 98% fo
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’Hazo tokana tsy mba ala’ One t
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Chapter 1 changes in resource avail
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Chapter 1 For analyses we narrowed
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Chapter 1 Fig. 2. Seasonal variatio
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Chapter 1 80 70 60 50 40 30 20 10 0
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Chapter 1 Fig 4. Comparison of data
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Chapter 1 Table 4. Data on phenolog
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Chapter 1 Relation with abiotic fac
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Chapter 1 Malagasy rainforests disp
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Chapter 1 Appendix I Continued Fami
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Chapter 1 Appendix I Continued Fami
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Malagasy proverb Drawing © Giusepp
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Chapter 2 the successful establishm
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Chapter 2 40 Table 1. The frugivore
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Chapter 2 Statistical analyses To v
Page 49 and 50: Chapter 2 Overall fruit trap data d
Page 51 and 52: Chapter 2 Table 3. Morphological an
Page 53 and 54: Chapter 2 depressed seeds (Fig. 3a)
Page 55 and 56: Chapter 2 Fig. 3. Ordination plot o
Page 57 and 58: Chapter 2 On a community level the
Page 59 and 60: Chapter 2 Appendix II. Plant-animal
Page 63 and 64: Malagasy proverb Lemur drawing © S
Page 65 and 66: Chapter 3 different consumer specie
Page 67 and 68: Chapter 3 Bollen et al. in press, C
Page 69 and 70: Chapter 3 64 Body 1,2 Body lenght (
Page 71 and 72: Chapter 3 dominant plant families s
Page 73 and 74: Chapter 3 Table 4. Morphological an
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Page 77 and 78: Chapter 3 Besides testing feeding s
Page 79 and 80: Chapter 3 and do not consume the fi
Page 81 and 82: Chapter 3 (1997) already pointed ou
Page 83 and 84: Chapter 3 1989; Galetti 1993; Saini
Page 85 and 86: Chapter 3 Appendix I. All plant spe
Page 87 and 88: Chapter 3 Other SD Hss Rats Rr Ew 8
Page 90 and 91: Pteropus rufus Feeding ecology of P
Page 92 and 93: Pteropus rufus number per species a
Page 94 and 95: Pteropus rufus Table 1 Continued Fr
Page 96 and 97: Pteropus rufus larger number of spe
Page 98 and 99: Table 2 Continued Number of seeds/d
Page 102 and 103: Pteropus rufus presence. On the oth
Page 104 and 105: Pteropus rufus Marshall (1983). Occ
Page 106 and 107: Coracopsis nigra The feeding ecolog
Page 108 and 109: Coracopsis nigra fruit pulp (70%) b
Page 110 and 111: Coracopsis nigra Table 1 Continued
Page 112 and 113: Coracopsis nigra Table 2. Morpholog
Page 115 and 116: Malagasy proverb Drawing of fruits
Page 117 and 118: Chapter 4 Most seed dispersal studi
Page 119 and 120: Chapter 4 Mean temperature is aroun
Page 121 and 122: Chapter 4 Number of seeds: 1-2, 3-1
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Page 129 and 130: Chapter 4 Table 5. Biochemical char
Page 131 and 132: Chapter 4 Chemistry At both sites,
Page 133 and 134: Chapter 4 considering species numbe
Page 137 and 138: ’Ny valala tsy indroa mandry eo a
Page 139 and 140: Chapter 5 environmental degradation
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Page 143 and 144: Chapter 5 Table 1. A list of plant
Page 145 and 146: Chapter 5 (Ganzhorn et al. 1999a).
Page 147 and 148: Chapter 5 Animals The degree of vul
Page 149 and 150: Chapter 5 comm.). Mixed species pla
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Chapter 5 data are needed we believ
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General conclusion as in other site
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General conclusion selection pressu
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General conclusion are consumed by
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Samenvatting omstandigheden zijn bo
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Samenvatting vruchtkenmerken. Beide
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Samenvatting zaadverspreiding moete
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Résumé température et précipita
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Résumé (Sud-est de Madagascar) et
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Résumé des frugivores sur l’év
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References Bollen A, Van Elsacker L
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References Dowsett JR (2000) Le sta
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References Godfrey LR, Junger WL, R
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References Janson CH, Chapman CA (1
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References Marshall AG (1983) Bats,
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References Rabevohitra R, Lowry PP,
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References Tattersall I (1982) The
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family Loganiaceae scientific name
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