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

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Chapter 3<br />

1989; Galetti 1993; Sa<strong>in</strong>i et al. 1994; Pizo et al. 1995; Corlett 1998; Renton 2001). For<br />

Madagascar Goodman et al. (1997a; 1997b) observed seed predation of Ficus seeds by<br />

Coracopsis spp., while Böhn<strong>in</strong>g Gaese et al. (1999) described C. nigra as an occasional<br />

seed disperser for Commiphora guillaum<strong>in</strong>i.<br />

Accord<strong>in</strong>g to the literature most Streptopelia spp. feed largely on dry seeds and fruits<br />

and destroy most of the seeds they swallow <strong>in</strong> their muscular gizzards (Corlett 1998).<br />

However, S. decaocto is said to regurgitate some large seeds (Corlett 1998). S. picturata<br />

was most often seen feed<strong>in</strong>g on the ground but no details on its feed<strong>in</strong>g behaviour could<br />

be obta<strong>in</strong>ed at our study site. Nevertheless this species is suspected to be a postdispersal<br />

seed predator destroy<strong>in</strong>g most seeds, which is confirmed by Goodman et al.<br />

(1997b). Obviously our list of food items sampled for this bird is largely underestimated<br />

compared to their actual diet.<br />

Before humans settled, the only seed predators <strong>in</strong> Sa<strong>in</strong>te Luce were Coracopsis spp.,<br />

S. picturata, and Eliurus webbi. The latter now has to contend also with a non-native<br />

rodent species (Rattus rattus) even though there are no <strong>in</strong>dications for food or habitat<br />

competition between these two rodent species (Ramanamanjato and Ganzhorn 2001;<br />

Ganzhorn 2003). Evidence of post-dispersal predation by rodents destroy<strong>in</strong>g the seeds of<br />

fifty plant species was found, but Goodman and Sterl<strong>in</strong>g (1996) and Ganzhorn et al.<br />

(1999a) suggested that native <strong>Malagasy</strong> rodents may store seeds <strong>in</strong> their burrows but no<br />

evidence has been found so far for Eliurus webbi <strong>in</strong> the <strong>littoral</strong> <strong>forest</strong>. Goodman (1994)<br />

describes a burrow of E. webbi <strong>in</strong> Andr<strong>in</strong>gitra conta<strong>in</strong><strong>in</strong>g 20 seeds of Cryptocarya sp. of<br />

which only half were eaten. In Sa<strong>in</strong>te Luce a few seeds were observed to escape total<br />

destruction when germ<strong>in</strong>at<strong>in</strong>g from the rodents’ food piles. This diet list can serve as a<br />

first <strong>in</strong>dication on the rodents’ food species <strong>in</strong> the <strong>littoral</strong> <strong>forest</strong>, tak<strong>in</strong>g <strong>in</strong>to account that<br />

completely digested seeds are miss<strong>in</strong>g here. Nevertheless by target<strong>in</strong>g seeds of selected<br />

species, rodents can significantly alter <strong>forest</strong> composition (DuPuy 1996; Spehn and<br />

Ganzhorn 2000).<br />

Besides the described <strong>frugivore</strong>s (Table 1), there are also other animal species <strong>in</strong><br />

Sa<strong>in</strong>te Luce that occasionally feed on fruits. Bird species such as Coua caerulea<br />

(Cuculidae) and Zosterops maderaspatana (Zosteropidae) (also observed by Goodman<br />

et al. 1997a) could be seen feed<strong>in</strong>g on fruits <strong>in</strong> Sa<strong>in</strong>te Luce, as well as the terrestrial<br />

mammal Tenrec ecaudatus (Tenrecidae) and the fruit bats Rousettus madagascariensis<br />

and Eidolon dupreanum (Pteropodidae), but the latter species were very difficult to track.<br />

Local people may also act as seed dispersers when collect<strong>in</strong>g several fruit species (n=12,<br />

App. I). As secondary seed dispersers Pheidole spp. (Myrmic<strong>in</strong>ae) seem to be important<br />

and they were observed to transport <strong>in</strong>tact seeds of at least 20 species to their ground<br />

nests, thus match<strong>in</strong>g the important role of ants <strong>in</strong> other <strong>forest</strong>s of Madagascar (Voigt et al.<br />

2002; App. I).<br />

As mentioned above, the dispersal quality of the <strong>frugivore</strong>s differs substantially.<br />

Frugivores are active at different times of the day and year, forage at different heights,<br />

have dist<strong>in</strong>ct feed<strong>in</strong>g behaviour, gut passage rates, and seed shadows. While birds may<br />

defecate from nearby or far away perches either with<strong>in</strong> primary <strong>forest</strong> or <strong>in</strong> the clear<strong>in</strong>gs,<br />

fly<strong>in</strong>g foxes defecate dur<strong>in</strong>g flight or under feed<strong>in</strong>g or sleep<strong>in</strong>g roosts, rodents<br />

concentrate seeds at burrows or feed<strong>in</strong>g sites and lemurs move seeds with<strong>in</strong> the <strong>forest</strong><br />

fragment only. In the end, the comb<strong>in</strong>ed action of a variety of fruit-eat<strong>in</strong>g vertebrates with<br />

dist<strong>in</strong>ct seed shadows produces a very heterogeneous transport of seeds, which is very<br />

important <strong>in</strong> the severely fragmented and degraded <strong>littoral</strong> <strong>forest</strong> to ensure regeneration<br />

of most plant species with<strong>in</strong> and outside <strong>forest</strong> fragments. So even though dietary overlap<br />

78

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