DOI: 10.5061/dryad.2ngf1vhkg
title: Data from: Fruit traits of pioneer trees structure seed dispersal across distances on tropical deforested landscapes: implications for restoration

description: Pioneer trees with fleshy fruits are typically planted in restoration
 projects to attract frugivores as a mean to increase dispersal and
 accelerate forest regeneration. However, differences in fruit traits of
 pioneer trees can potentially influence dispersal and their restoration
 outcomes. Here we investigated the effects of bird and plant traits, and
 distance to forest fragments, on the seed rain using a tree-planting
 experiment replicated in 12 deforested sites in Brazil. Factors were fruit
 traits of pioneer trees (wind-dispersed, bird-dispersed with lipids or
 with carbohydrates, and controls) and distance (10, 50, 300 m) from forest
 fragments. We found that density and richness of birds and seeds decreased
 exponentially with distance from fragments, yet these effects were minor
 compared to the effects of fruit traits on the structure of the seed rain.
 Overall, plots with fleshy-fruited pioneers attracted much greater bird
 activity and seed dispersal than plots with wind-dispersal pioneers and
 the controls. For instance, plots with carbohydrate-rich fruits received
 more than twice the average species richness and density of birds and
 seeds of plots with lipid-rich pioneer trees, surpassing wind-dispersed
 pioneers by more than 80%, and controls by over 90%. Furthermore, the
 fruit trait treatments resulted in morphological shifts in the average
 traits of visiting birds. Significant differences in bill gape and flight
 capacities (wing-loading) were associated with the differences in the seed
 rain associated with each treatments. Synthesis and applications.
 Understanding how trait-matching processes mediating mutualistic seed
 dispersal by frugivores interact with distance-dependent dispersal
 limitation on deforested tropical landscapes is critical for improving
 forest restoration efforts. This is especially relevant in the context of
 applied nucleation. As shown here, avian seed dispersal can thus be
 manipulated in restoration projects in order to increase connectivity and
 speed up forest recovery and the provision of the multiple ecosystem
 services that follow forest succession.
description: <i>Study sites</i> We conducted
 the experiment in 12 sites in Paranapanema, São Paulo, Brazil
 (23°23'S, 48°43'W, Fig. 1). Paranapanema is located at ca. 600 m
 a.s.l. on the watershed region of the Alto Paranapanema river (Cielo-Filho
 et al., 2009). Average precipitation is 1.407,9 mm mostly during
 December-March, and a mean temperature of 18 °C (Cielo-Filho et al.,
 2009). Atlantic forest fragments cover ~6% of the Paranapanema landscape
 (Fundação SOS Mata Atlântica, 2013). Our study sites were located on
 private cattle pastures and fragments of primary and secondary
 semideciduous forest &gt;30 years old and 12.2-98.8 ha in
 size.   <i>Experiment set
 up</i> Between December 2016 and November 2018 we
 conducted a full-factorial experiment using two explanatory variables:
 <i>distance</i> to forest fragment at three levels: 10, 50,
 300 m, and <i>fruit traits </i>of pioneer trees<i>
 </i>at three levels: <i>Heliocarpus popayanensis</i>
 Kunth with wind-dispersed fruits, <i>Acnistus arborescens</i>
 Schltdl, with bird-dispersed fruits, and <i>Trema
 micrantha</i> (L.) Blume also with bird-dispersed fruits, plus a
 control treatment without any tree. Thus, the 4 x 3 design yielded 12
 types of plots that were replicated once in each of 12 localities in
 Paranapanema, totaling 144 (Fig. 1). The relative position of each
 <i>Fruit trait </i>treatment<i> </i>plot was
 randomly determined within each distance class, and spaced the others by a
 distance equal to their distance to the from fragments. The two
 bird-dispersed trees (<i>Acnistus</i> &
 <i>Trema</i>) differed in their fruit traits:
 <i>Acnistus</i> has larger carbohydrate-rich fruits, while
 <i>Trema</i> has small lipid-rich fruits (See Table S1). Plots
 measured 4.5 x 4.5 meters and were fenced with barbwire to keep cattle out
 (Fig. S1). Pioneer trees from all species had similar height, crown, and
 fruit crop (Tables S1-S2) and were grown nurseries until a height of ~1.5
 m. A single tree was planted at the center of plots requiring trees, and
 fertilized with 300 g of 15-18-28 (NPK) every six months. Grasses were
 mechanically controlled during the entire experiment.
   <i>Bird activity</i>
 We recorded bird visits in all experimental plots using a
 combination of video recording and focal observation with effort equally
 distributed among plots. We used 28 video cameras from four different
 brands (ten Bushnell Trophy Cam, six Bushnell Trophy Cam HD, six Tigrinus,
 and six GoPro Hero 3). Camera traps operated 24 h/day recording 88,714.7
 h, while the GoPro Hero 3 cameras filmed for 2 hours per day for, 288 h.
 To prevent sampling bias associated with camera models, all units were
 systematically rotated to sample for the same amount of time in each
 experimental plot across the twelve study sites. In addition, in each plot
 one observer (P. Camargo) recorded bird visits using a pair of binoculars
 from a distance of 50m for 20 minutes, during morning hours (0730-1030),
 once a month, for 5 months.  
 <i>Seed rain</i> We sampled the
 seed rain of all plots using one 0.25-m² seed trap lined with a 0.2-mm
 nylon mesh and located at the center of the enclosure and covered with a
 2.5 x 2.5 cm wire mesh (Fig. S1). The sticky paste
 Formifuu<sup>®</sup> was applied to the support posts of traps
 to exclude ants. Traps were collected each month and seeds counted and
 identified with the aid of a dissecting scope and reference books and seed
 collections. Seeds from grasses or from the species of the pioneer tree
 species in a plot were excluded from analyses.
   <i>Bird and seed
 traits</i> Bird traits included % fruit in diet
 (Wilman et al. 2014), wing-loading as a proxy of bird movement
 capacity, and gape-width as a proxy to their upper seed size dispersal
 potential. Wing-loading was calculated as:
 <i>WLo= </i><i>BM</i><i>2×WL</i> where BM is the bird body mass and WL is wing length (species measurements were obtained from Rodrigues et al. 2019). As loading value increases, the movement capacity of a bird species decreases. We classified dispersed seeds by dispersal mode: bird-dispersed, wind-dispersed, and gravity-dispersed. In addition, we calculated seed mass and approximated lipid content of fruit in a rank scale (1 - 0 to 10% lipids, 2 - 10 to 20%, 3 - 20 to 30% and 4 - above 30%) adapted from Bello et al. (2017).