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Quantity component of the effectiveness of seed dispersal by birds in ...

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BOSQUE 32(1): 39-45, 2011Seed <strong>dispersal</strong> <strong>by</strong> <strong>birds</strong> <strong>in</strong> <strong>the</strong> forest <strong>of</strong> Chiloéperiod, as has been recorded <strong>in</strong> o<strong>the</strong>r different temperatera<strong>in</strong>forests <strong>in</strong> sou<strong>the</strong>rn Chile dur<strong>in</strong>g autumn and w<strong>in</strong>termonths (Sabag 1993, Hernández 1995).Although this study does not prove that T. falcklandii is alegitimate disperser for <strong>the</strong> tree species studied, <strong>the</strong> evidence 2susta<strong>in</strong>s that this species may well be a disperser <strong>of</strong> viable<strong>seed</strong>s <strong>of</strong> both L. apiculata and A. punctatum (Figueroa andCastro 2003). Seed germ<strong>in</strong>ation <strong>of</strong> both L. apiculata andA. punctatum obta<strong>in</strong>ed from bird feces collected at random<strong>in</strong> <strong>the</strong> field, dur<strong>in</strong>g <strong>the</strong> autumn, is similar to <strong>the</strong> <strong>seed</strong>germ<strong>in</strong>ation obta<strong>in</strong>ed <strong>in</strong> fruits recently collected on <strong>the</strong>mo<strong>the</strong>r plants (Figueroa and Castro 2003). Accord<strong>in</strong>g toour results <strong>of</strong> bird abundance, <strong>the</strong>re is a high probabilitythat <strong>the</strong> feces collected at random <strong>in</strong> <strong>the</strong> field dur<strong>in</strong>gautumn belong to T. falcklandii. However, <strong>the</strong> hypo<strong>the</strong>sisshould be corroborated <strong>in</strong> later studies.These results confirm that temperate ra<strong>in</strong>forests <strong>of</strong>South America have low diversity <strong>of</strong> frugivorous <strong>birds</strong>(Sabag 1993, Hernández 1995, Amico and Aizen 2005),which contrasts with <strong>the</strong> observations carried out <strong>in</strong> o<strong>the</strong>rtemperate regions around <strong>the</strong> world (Stepanian 1982, Debusscheand Isenman 1989, Stanley and Lill 2002, Jordanoet al. 2007). This is most likely due to <strong>the</strong>ir isolation fromo<strong>the</strong>r South America forests s<strong>in</strong>ce <strong>the</strong> Pliocene (H<strong>in</strong>ojosaand Villagrán 1997).Specifically, <strong>the</strong> “olivillo” forest studied has <strong>the</strong> lowestdiversity <strong>of</strong> frugivorous <strong>birds</strong>. Nearly all <strong>of</strong> <strong>the</strong> feed<strong>in</strong>gvisits on A. punctatum trees were <strong>of</strong> T. falcklandii and< 5 % were C. araucana. In addition, T. falcklandii is also<strong>the</strong> ma<strong>in</strong> visitor <strong>of</strong> L. apiculata, with 65 % <strong>of</strong> feed<strong>in</strong>g visits.Never<strong>the</strong>less, o<strong>the</strong>r three bird species are responsiblefor <strong>the</strong> rema<strong>in</strong><strong>in</strong>g L. apiculata fruit consumption.In <strong>the</strong> study period, T. falcklandii was sighted 18 timesmore frequently <strong>in</strong> <strong>the</strong> “olivillo” patch than <strong>in</strong> <strong>the</strong> secondaryforest where L. apiculata was common. However,T. falcklandii has similar rates <strong>of</strong> feed<strong>in</strong>g visits to both treespecies (2.2 and 1.8 bird visits per hours <strong>in</strong> L. apiculataand A. punctatum, respectively). Regard<strong>in</strong>g <strong>the</strong> dispersed<strong>seed</strong>s per hour (<strong>the</strong> quantitative <strong>component</strong> <strong>of</strong> <strong>effectiveness</strong>),<strong>in</strong> this study T. falcklandii was less effective onA. punctatum fruit (3.4 <strong>seed</strong>s per hours) than when feed<strong>in</strong>gon L. apiculata fruit (49.2 <strong>seed</strong>s per hours). Also, <strong>in</strong>this last tree, <strong>the</strong> quantity <strong>of</strong> <strong>seed</strong>s dispersed per hour wassignificantly greater for T. falcklandii, <strong>in</strong>dicat<strong>in</strong>g probablyhigher <strong>effectiveness</strong> <strong>of</strong> <strong>the</strong> quantitative <strong>component</strong> dueto greater abundance <strong>in</strong> <strong>the</strong> study site and more <strong>seed</strong>s perfruit. The rema<strong>in</strong><strong>in</strong>g bird species dispersed < 30 % <strong>of</strong> <strong>the</strong>L. apiculata <strong>seed</strong>s. This greater abundance <strong>of</strong> T. falcklandii,recorded dur<strong>in</strong>g <strong>the</strong> study period <strong>in</strong> Chiloé, may showthat <strong>the</strong> number <strong>of</strong> visits a disperser makes appears to be abetter predictor <strong>of</strong> <strong>the</strong> total quantity <strong>of</strong> <strong>seed</strong> dispersed than<strong>the</strong> number <strong>of</strong> fruits <strong>in</strong>gested per visit (Schupp 1993).In addition, a study 3 directly done on <strong>the</strong> “olivillo” understoryshows that > 90 % <strong>of</strong> <strong>the</strong> A. punctatum propagules2M Salvande: data not published.3M Salvande: data not published.were <strong>in</strong>tact <strong>in</strong> <strong>the</strong> site and would mean that propagules fallfrom trees without be<strong>in</strong>g consumed <strong>by</strong> frugivores and/orthat animals handle <strong>the</strong>m <strong>in</strong> a deficient way (Snow andSnow 1988). Studies <strong>in</strong> different communities also haveshown crops that are ignored <strong>by</strong> endozoochorous dispersers(Snow and Snow 1988, Fuentes 2000). In our study,<strong>the</strong> proportion <strong>of</strong> A. punctatum fruits dispersed <strong>by</strong> <strong>birds</strong>,as well as <strong>the</strong> number <strong>of</strong> fruits <strong>in</strong>gested per visit, is verylow when compared to data found for L. apiculata ando<strong>the</strong>r woody species (Stanley and Lill 2002, Obeso andFernández-Calvo 2002). Our results seem to suggest thatfruit size would be a limit<strong>in</strong>g factor for <strong>birds</strong> forag<strong>in</strong>g onA. punctatum fruits. L. apiculata fruits are smaller <strong>in</strong> sizethan those <strong>of</strong> A. punctatum (table 1). Therefore, <strong>in</strong> <strong>the</strong> temperatera<strong>in</strong>forests <strong>of</strong> sou<strong>the</strong>rn Chile, L. apiculata could bea less discrim<strong>in</strong>atory source <strong>of</strong> ripe fruits than A. punctatum,because <strong>the</strong> bird species present <strong>in</strong> <strong>the</strong> region generallyhave beak gapes smaller than A. punctatum fruit diameter(Armesto et al. 1987). This may also expla<strong>in</strong> why <strong>the</strong>lowest diversity <strong>of</strong> frugivorous <strong>birds</strong> is found <strong>in</strong> <strong>the</strong> patchdom<strong>in</strong>ated <strong>by</strong> A. punctatum and <strong>the</strong> low <strong>effectiveness</strong> <strong>of</strong>T. falcklandii on A. punctatum fruits, at least with regardto <strong>the</strong> quantitative <strong>component</strong>s <strong>of</strong> disperser <strong>effectiveness</strong>.In addition, <strong>the</strong> relative amount <strong>of</strong> pulp per fruit is lower <strong>in</strong>A. punctatum than <strong>in</strong> L. apiculata. However, we do nothave data regard<strong>in</strong>g nutrient content relative to <strong>the</strong> pulp forboth species, which are important variables when consider<strong>in</strong>gfruits as food resources for frugivores (Jordano 1995).In conclusion, <strong>in</strong> <strong>the</strong> study site we observed fourfrugivorous <strong>birds</strong> dur<strong>in</strong>g autumn-w<strong>in</strong>ter (T. falcklandii,P. rara, C. pitius and C. curaeus). Similarly, <strong>the</strong>re is evidencethat T. falcklandii is an important <strong>seed</strong> disperserdur<strong>in</strong>g autumn-w<strong>in</strong>ter <strong>in</strong> <strong>the</strong> temperate forests <strong>of</strong> SouthAmerica (Amico and Aizen 2005). Regard<strong>in</strong>g quantity<strong>component</strong> <strong>of</strong> disperser <strong>effectiveness</strong>, this study showsthat T. falcklandii would be <strong>the</strong> most effective disperserbird <strong>in</strong> <strong>the</strong> ra<strong>in</strong>forests <strong>of</strong> Chiloé. First, fleshy fruits produceddur<strong>in</strong>g austral autumn and w<strong>in</strong>ter were pr<strong>in</strong>cipally<strong>in</strong>gested and <strong>the</strong> <strong>seed</strong>s probably dispersed <strong>by</strong> T. falcklandii.Secondly, P. rara, which consumes a great number <strong>of</strong>L. apiculata fruits per visit, is present <strong>in</strong> lower abundanceand disperses a lower number <strong>of</strong> <strong>seed</strong>s dur<strong>in</strong>g this period.F<strong>in</strong>ally, as <strong>the</strong> measurements were obta<strong>in</strong>ed on quantitative<strong>component</strong>s <strong>of</strong> disperser <strong>effectiveness</strong> and only <strong>in</strong>one study site <strong>by</strong> tree species, additional <strong>in</strong>vestigation isrecommended to extend <strong>the</strong> research over qualitative <strong>component</strong>s<strong>of</strong> <strong>effectiveness</strong>, and over <strong>the</strong> spatial and <strong>in</strong>terannualvariation <strong>of</strong> <strong>the</strong> Chiloé forests.ACKNOWLEDGEMENTSDur<strong>in</strong>g his stay <strong>in</strong> Chile, M Salvande was supported <strong>by</strong>a pre-doctoral degree granted <strong>by</strong> <strong>the</strong> Spanish M<strong>in</strong>istry <strong>of</strong>Science and Education (AP97-33299901), and he wishesto thank all <strong>the</strong> staff <strong>in</strong> “Senda Darw<strong>in</strong>” for <strong>the</strong>ir warmassistance dur<strong>in</strong>g his stay<strong>in</strong>g <strong>in</strong> Chiloé. JA Figueroa was43

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