Fruit-frugivore interactions in a Malagasy littoral forest - Universiteit ...

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Coracopsis nigra Table 2. Morphological variables within the diet of Coracopsis nigra , Streptopelia picturata and the rodents (including E. webbi and Rattus rattus ) are shown. A comparison is made for the continuous variables through Kruskal Wallis test and for the class variables through contingency tables. C. nigra S. picturata rodents Kruskal Wallis median (N=37) median (N=13) median (N=50) H df P Number of seeds 1 1 1 0.13 2 0.93 Fruit length 12.28 9.55 19.04 22.35 2
Chapter 3b behaviour of parrots worldwide, many authors agree on their role as seed predators (Janzen 1981; Jordano 1983; Clout 1989; Galetti 1993; Saini et al. 1994; Pizo et al. 1995; Corlett 1998; Renton 2001). C. nigra and possibly also C. vasa can certainly be considered as granivores in Sainte Luce rather than frugivores, as they are so often referred to in literature. As such they vary from other seed predators in many different aspects. C. nigra is a pre-dispersal seed predator, which feeds in the tree crown only during daylight on both ripe and unripe fruits, which are medium in size. It can further tolerate high tannin contents and eats non-zoochorous fruits as well without any fruit pulp at all. S. picturata is a post-dispersal seed predator which eats small seeds of juicy fruits on the ground only, either with a protective seed coat or not. The rodents both feed on the ground as in the tree at night, but most often carry seeds away to a feeding site. They eat the largest seeds of mostly drupes and may occasionally be involved in secondary seed dispersal as well. Nevertheless they can be mainly considered as post-dispersal seed predators for the majority of consumed fruit species. As shown in Figure 1 C. nigra has an almost constant year-round food availability as unripe fruits in particular stay long periods on the tree before ripening. On the other hand seeds stay longer periods available on the ground than in the tree, which is favourable for both the rodents and S. picturata. So, apparently there is a good niche separation among all seed predators in this ecosystem, which is mainly based on fruit and seed size, feeding height and activity pattern. In conclusion, C. nigra destroys seeds of numerous endemic plant species in Sainte Luce and should therefore be considered a seed predator in this ecosystem. In general, they split and thus destroy the seed with their bill, eat it and reject the surrounding pulp. Seed dispersal does occasionally occur when parrots fly away from the parent plant, dropping several intact fruits or the remains of soft and juicy berries. Zoochorous tree species have evolved phenological, morphological and biochemical flower, fruit and seed characteristics to attract efficient pollinators and seed dispersers, which can assure successful plant regeneration. However, the behaviour of granivorous animals offers the plant no apparent advantage. Nevertheless, most food species consumed by these parrots are also eaten by other frugivores, which act as seed dispersers. Dietary overlap among granivores is limited and the different seed predators seem to occupy separate trophical niches in this ecosystem. ACKNOWLEDGEMENTS We would like to thank QMM (QIT Madagascar Minerals) for providing logistics and infrastructure at the study site. We are very grateful to Steve Goodman and Jörg Ganzhorn for their extensive revisions and useful suggestions. This study was carried out under a Collaboration Agreement between the Department of Animal Biology and the Department of Anthropology of the University of Antananarivo, the Institute of Zoology of Hamburg University and QMM. The first author was supported by a grant from the Belgian Fund for Scientific Research, Flanders (FWO). We thank the Flemish Government for its structural support for the Centre of Research and Conservation (CRC) of the Royal Zoological Society of Antwerp (RZSA). 108
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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
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Chapter 2 Overall fruit trap data d
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Chapter 2 Table 3. Morphological an
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Chapter 2 depressed seeds (Fig. 3a)
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Chapter 2 Fig. 3. Ordination plot o
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Chapter 2 On a community level the
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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
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Page 75 and 76: Chapter 3 Table 4 Continued Paramet
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 100 and 101: Pteropus rufus Food selection: expl
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 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
Page 123 and 124: Chapter 4 Fig. 2. Monthly fruit ava
Page 125 and 126: Chapter 4 120 Table 3. Soil charact
Page 127 and 128: Chapter 4 Table 4. Morphological ch
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
Page 141 and 142: Chapter 5 at a very low density in
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
Page 151 and 152: Chapter 5 data are needed we believ
Page 153 and 154: General conclusion as in other site
Page 155 and 156: General conclusion selection pressu
Page 157 and 158: General conclusion are consumed by
Page 159 and 160: Samenvatting omstandigheden zijn bo
Page 161 and 162: 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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