Consumption of Euterpe edulis fruit by wildlife: implications for conservation and management of the Southern Brazilian Atlantic Forest

Abstract

This study aimed to measure the wildlife consumption of Euterpe edulis fruit and use this data to discuss management possibilities. To estimate infructescence fruit volume consumed, collectors were installed in fruit-bearing palms. To characterize consumption from the ground, samples were placed next to fruiting palms. To identify wildlife and their activities, camera traps were installed in infructescences and on the ground. The results suggested that there was a small fruit surplus (1.8 %), and this finding indicated the possibility of a harvest to reduce food for the wildlife. However, recurrent variations in the annual fruit production (21.4 %) were also noted, and suggested that wildlife could tolerate some fruit harvesting. Thus, a harvest could be restricted to fruit volume that exceeds the annual average (94 kg/ha/year). Turdus flavipes, a migratory bird, was the most active species in the dispersal of seeds; this finding indicates the need for broader conservation strategies. Wildlife composition also changed along with the fruiting, and this alteration suggests that dependence on the fruit is variable among different species. Seed germination and seedling mortality were high, results that indicate that local conditions may have a predominant effect on seed volume in natural regeneration density.

Full text

Anais da Academia Brasileira de Ciências (2019) 91(1): e20180537
(Annals of the Brazilian Academy of Sciences)
Printed version ISSN 0001-3765 / Online version ISSN 1678-2690
http://dx.doi.org/10.1590/0001-3765201920180537
www.scielo.br/aabc | www.fb.com/aabcjournal
An Acad Bras Cienc (2019) 91(1)
BIOLOGICAL SCIENCES
Consumption of Euterpe edulis fruit by wildlife: implications for conservation
and management of the Southern Brazilian Atlantic Forest
JULIANO Z. DA SILVA and MAURÍCIO S. DOS REIS
Universidade Federal de Santa Catarina, Centro de Ciências Agrárias, Departamento de Fitotecnia,
Rodovia Admar Gonzaga, 1346, Itacorubi, 88034-001 Florianópolis, SC, Brazil
Manuscript received on May 29, 2018; accepted for publication on July 16, 2018
How to cite: SILVA JZ AND REIS MS. 2019. Consumption of Euterpe edulis fruit by wildlife: implications for
conservation and management of the Southern Brazilian Atlantic Forest. An Acad Bras Cienc 91: e20180537. DOI
10.1590/0001-3765201920180537.
Abstract: This study aimed to measure the wildlife consumption of Euterpe edulis fruit and use this data
to discuss management possibilities. To estimate infructescence fruit volume consumed, collectors were
installed in fruit-bearing palms. To characterize consumption from the ground, samples were placed next to
fruiting palms. To identify wildlife and their activities, camera traps were installed in infructescences and
on the ground. The results suggested that there was a small fruit surplus (1.8 %), and this nding indicated
the possibility of a harvest to reduce food for the wildlife. However, recurrent variations in the annual
fruit production (21.4 %) were also noted, and suggested that wildlife could tolerate some fruit harvesting.
Thus, a harvest could be restricted to fruit volume that exceeds the annual average (94 kg/ha/year). Turdus
avipes, a migratory bird, was the most active species in the dispersal of seeds; this nding indicates the
need for broader conservation strategies. Wildlife composition also changed along with the fruiting, and
this alteration suggests that dependence on the fruit is variable among dierent species. Seed germination
and seedling mortality were high, results that indicate that local conditions may have a predominant eect
on seed volume in natural regeneration density.
Key words: natural regeneration, non-timber forest products, rain forest, seed dispersal, seed predation.
Correspondence to: Juliano Zagos da Silva
E-mail: jzagos@yahoo.com.br
ORCid: http://orcid.org/0000-0001-6296-0814
INTRODUCTION
Many palms produce fruit of great economic
importance. In South America, Bernal et al.
(2011) described the human use of 96 palm
species, of which the major producers of fruit are:
Bactris gasipaes, Mauritia exuosa, Oenocarpus
bataua, Euterpe oleracea, Euterpe precatoria,
Astrocaryum aculeatum, Acrocomia aculeata,
Aiphanes horrida, and Bactris guineensis. Palm
and g trees are commonly cited as key tropical
species and are hyperdominant as food sources for
wildlife (Terborgh 1986, Lambert and Marshall
1991, Kinnaird 1992, Peres 1994, Fredericksen
et al. 1999, Galetti et al. 1999, Genini et al. 2009,
Staggemeier et al. 2017). However, competition
between humans and animals for these resources,
coupled with an actual reduction in the resources
caused by deforestation and forest fragmentation,
is a growing reality, especially in the Brazilian
Atlantic Forest, the primary location of Euterpe
edulis.

JULIANO Z. DA SILVA and MAURÍCIO S. DOS REIS CONSUMPTION OF Euterpe edulis FRUIT BY WILDLIFE
An Acad Bras Cienc (2019) 91(1) e20180537 2 | 20
In many cases, reduction and change in
wildlife composition have been related to reduced
availability of fruit within forests (Kinnaird 1992,
Galetti and Aleixo 1998, Moegenburg and Levey
2002, 2003, Weterings et al. 2008, Bicknell and
Peres 2010, Galetti et al. 2013), particularly if
coupled with destructive extractive methods, such
as cutting plants to collect fruit (Bernal et al. 2011).
Additionally, studies related to the sustainable
management of non-timber forest products
(NTFPs) are fundamentally concerned with natural
regeneration and demographic growth of the target
species (Pinard and Putz 1992, Godoy and Bawa
1993, Silva Matos et al. 1999, Reis et al. 2000,
Freckleton et al. 2003, Portela et al. 2010, Pandey
and Shackleton 2012), as well as the maintenance
of genetic diversity (Reis 1996, Conte et al. 2003,
Seoane et al. 2005, Silva and Reis 2010). Only a
few studies, such as Fredericksen et al. (1999),
concern the availability of resources for wildlife.
The sustainability perspective normally places
wildlife needs in second place, especially when
it concerns the consumption of fruit pulp without
seed destruction. Due to experimental diculties,
or simply because of the intrinsic idea that
animals may collect and consume other resources,
the proportion of fruit from a given species that
should remain in the forest to feed wildlife has
been ignored. However, the question, “What are
the consequences of disregarding or ignoring the
fruiting of species considered key in the forest
environment?”, is of great importance, especially
when a specic fruiting period is related to a period
of resource scarcity.
E. edulis has been the main producer of heart of
palm in the Brazilian Atlantic Forest since the 1960s
when it became the target of intensive commercial
exploitation (Reis and Reis 2000). Aside from
serving as raw material for the production of heart
of palm, the fruit from this species is currently
used in Southern and Southeastern Brazil as an
alternative to açaí (Euterpe oleracea and Euterpe
precatoria) (Barroso et al. 2010, Favreto et al.
2010, Trevisan et al. 2015); the pulp (exocarp and
mesocarp) is extracted as a high-energy food.
E. edulis fruit extraction has not been
intensively exploited and is a less aggressive and
more sustainable possibility, since extraction does
not require removing the individual palm from
the forest. In addition, economic value, market
acceptance, and consumer demand for this product
have all increased signicantly in Southern Brazil
(Trevisan et al. 2015). Thus, interest in developing
research and standards to support this new supply
chain has grown on the part of both farmers and
government agencies (Barroso et al. 2010, Favreto
et al. 2010, Justen et al. 2012, Trevisan et al.
2015). On the other hand, the fruit from this palm,
especially when mature, are an important food for
wildlife, and its supply usually occurs between 4
and 6 months of the year (Mantovani and Morellato
2000, Galetti et al. 1999, Genini et al. 2009,
Staggemeier et al. 2017, Silva and Reis 2018). A
total of 58 bird and 21 mammal species are known
to use this resource, and some of these species are
considered endangered (Galetti and Chivers 1995,
Galetti and Aleixo 1998, Galetti et al. 2000, 2001,
Reis and Kageyama 2000, Mikich 2002, Pizo et
al. 2002, Galetti et al. 2013). Thus, E. edulis is
considered a hyperdominant and keystone species
within the Atlantic Forest ecosystem (Galetti and
Chivers 1995, Galetti et al. 2000, 2001, Mikich
2002, Pizo et al. 2002, Staggemeier et al. 2017).
Studies on E. edulis include: Galetti and
Chivers (1995), Reis (1996), Galetti and Aleixo
(1998), Conte et al. (2000), Galetti et al. (2000),
Mantovani and Morellato (2000), Reis and
Kageyama (2000), Freckleton et al. (2003), Fantini
and Guries (2007), Genini et al. (2009), Barroso
et al. (2010), Favreto et al. (2010), Portela et al.
(2010), Silva and Reis (2010), and Paludo et al.
(2012). These authors have reported on fruit
production, consumption, and extraction, but
have neglected fruit consumption by wildlife,

JULIANO Z. DA SILVA and MAURÍCIO S. DOS REIS CONSUMPTION OF Euterpe edulis FRUIT BY WILDLIFE
An Acad Bras Cienc (2019) 91(1) e20180537 3 | 20
especially from the perspective of supporting
sustainable extraction of the fruit. Such studies are
generally directed toward the behavior of wildlife,
the dynamics of regeneration, or reproductive
phenology and maintenance of genetic diversity.
Thus, this paper aims to broaden the study of E.
edulis by examining fruit consumption by wildlife,
and looks to establish quantifiable surpluses by
identifying the percentage of fruit consumed and
the percentage of seeds that remain viable for
germination. We hypothesize that such surpluses
can serve as the basis for sustainable management
strategies and thereby maintain interactions with
wildlife and natural regeneration dynamics.
MATERIALS AND METHODS
STUDY AREA
The study site was located in the Ibirama National
Forest (Ibirama FLONA), Santa Catarina, Brazil.
This area contains 463 ha of native forest that is
connected to other larger forest fragments (http://
www4. icmbio.gov.br/flonaibirama). The local
vegetation is composed of montane Atlantic
rainforest (Klein et al. 1986) and exhibits structural
characteristics similar to areas of primary forest
cover.
QUANTIFICATION OF FRUIT PRODUCTION
To characterize reproductive palm density, the
number of seedlings in class I (with height of
insertion of the youngest leaf less than 10 cm), and
fruit production per area, data were obtained from
24 existing permanent plots (40 x 40 m) (Conte
et al. 2000) for which a phenological assessment
was carried out for all E. edulis individuals (888
reproductive palms) during the 2 years of evaluation
(2008 and 2009). In order to characterize annual
variation in the number of reproductive palms,
plants that emitted inorescences were recorded
in these same plots for 11 fruiting seasons (2004 to
2009, 2011, and 2013 to 2016).
FRUIT AND SEED USE, AND GERMINATION
PERCENTAGE
To quantify fruit use in infructescences and the
number of viable seeds for germination, 114
collectors (Supplementary Material, Figure S1a)
were installed, one for each palm infructescence on
92 reproductive palms. This installation allowed us
to identify and quantify predation, transport, and
pulping.
After all fruit was counted on each
infructescence, the collectors were installed below
fruit bunches by xing them on the stipe of palms
(Figure S1a). Each collector consisted of an iron
ring with a 3 m circumference and a sewn funnel-
shaped screen net. Of the 114 collectors, 80 were
positioned in 2008 and 34 in 2009 in order to
sample dierent reproductive cycles.
All fallen fruit in the collectors were visually
assessed and quantied for signs of predation (seed
damage due to insect drills, vertebrates, or fungi),
pulping (pulp exocarp and mesocarp removal
without damage to the seed), cracks (pulp and
endosperm cracks through seed growth), dryness
(endosperm was absent and no exocarp damages),
scratches (marks of nails, beaks, or teeth, but
without pulping or predation; these observations
characterize fruit loss during feeding), or whether it
remained intact. During each evaluation of the fallen
fruit in the collectors, 20 fruit per infructescence
were randomly sampled and cut. This assessment
aimed to identify the presence of seed damage that
was not externally perceptible (caused by insects
or fungi) and to correct the number of intact fruit in
each evaluation.
Fruit that did not fall into the collectors were
considered dispersed or transported primarily by
wildlife. This conclusion was based on results
obtained with the installation of camera traps,
as described below. The volume of primary seed
dispersal was corrected using the percentage
of predation observed in the infructescences.

JULIANO Z. DA SILVA and MAURÍCIO S. DOS REIS CONSUMPTION OF Euterpe edulis FRUIT BY WILDLIFE
An Acad Bras Cienc (2019) 91(1) e20180537 4 | 20
The percentage of pulped fruit (dispersers are
regurgitants or seed defecators; see Galetti et al.
2013) was also corrected for the rates of predation,
rot (when it was not possible to identify the agent;
most likely fungi or bacteria), secondary transport,
and any intact remnants, as observed in seed
samples arranged on the soil (609 samples from
50,065 total seeds).
To study the activities of wildlife toward fruit
on the ground, samples of unripe (47 samples
from 2,551 total fruit), ripe (251 samples from
11,244 total fruit), dried (346 samples from 9,269
total fruit), and pulped fruit (609 samples from
50,065 total seeds) were placed on the ground
below collectors for each reproductive palm and
evaluated during 2008 and 2009. The samples were
placed in dened spaces where fruit were separated
by type. Evaluation and replacement of fruit were
performed weekly, and the intensity was determined
by the amount of fruit that fell into the collectors. In
these samples, fruit and seeds were also evaluated
for pulping, transport, predation, decay, percent
that remained intact, and seedling mortality that
originated from seed germination. Due to fruit or
seed replacement on soil samples, the germination
percentage was evaluated in a greenhouse. Thus,
we used 8,222 seeds pulped by animals from 67
palm with installed collectors.
When fruit or seeds disappeared from soil
samples, these were considered to be secondary
dispersal or transport activities, and these volumes
were later corrected for percentages of predation,
rot, and quantity that remained intact. Corrections
in these cases depended on the object of study. Thus,
for seeds, only one correction was performed, using
data from seed samples arranged on the soil (609
samples from 50,065 total seeds); two corrections
were required for fruit: one based on samples of
fruit arranged on the soil (251 samples from 11,244
total fruit) and another based on seeds (609 samples
from 50,065 total seeds). Double correction of
the fruit transported at soil level was performed
based on the continuity of wildlife activities on the
volume of pulped fruit, which behave like seeds
when pulp is lost. Between 2013 and 2014, camera
traps were also installed to monitor seeds and fruit
arranged at ground level, as described below.
CHARACTERIZATION OF WILDLIFE SPECIES AND
THEIR ACTIVITIES
We installed camera traps to identify wildlife
species and their interaction frequencies, and
quantify the number of fruit and seeds dispersed,
predated, pulped, and regurgitated or defecated
(for seeds) in infructescences or at the base of the
reproductive palms. Monitoring was performed
monthly, including periods of 15 to 20 days per
month per trap where images were recorded day
and night (24 h) throughout the entire fruiting
period that occurred in 2013 (7 months) and during
4 months of fruiting that occurred in 2014. The
camera conguration used was: high sensitivity to
movement, videos up to 1 min, and 2 min interval
between recordings.
Up to 11 camera traps per month were installed
to monitor infructescences of 64 reproductive palms:
34 in 2013 and 30 in 2014 (the same plants where
the collectors were installed). Trap installation
was either directly among the leaf sheaths of the
fruiting plant (Fig. S1b) or indirectly on the stipe of
nearby plants and not more than 2 m away from the
monitored infructescence (Fig. S1c).
To monitor fruit and seeds arranged on the soil,
up to 6 camera traps per month and samples with
fruit and seeds (with monthly replenishment) were
used. The samples were kept separate and distant
at a maximum of 1.5 m from the traps. Traps were
installed on the same plants where the collectors
were mounted. They were xed at 50 cm in height
on 30 fruiting palms (Fig. S1d): 19 in 2013 and 11
in 2014.
The study area was also evaluated for the
average size of seeds regurgitated by frugivores and
the richness of birds classied as follows: frugivory

JULIANO Z. DA SILVA and MAURÍCIO S. DOS REIS CONSUMPTION OF Euterpe edulis FRUIT BY WILDLIFE
An Acad Bras Cienc (2019) 91(1) e20180537 5 | 20
score = 1: sporadic, 2: moderate, and 3: extensive
frugivory; feeding behavior: R: regurgitate seeds,
D: defecate seeds, SP: seed predator, and PC:
pulp consumer; classication follows the method
proposed by Galetti et al. (2013). The objective of
this procedure was to compare the study area with
areas considered defaunate and non-defaunate as
studied by Galetti et al. (2013). The regurgitated
seeds were collected during fruiting from 2013 to
2017, with a minimum distance of 15 m between
seeds. A total of 3,772 seeds were measured in
three axes. We used the smallest measure obtained
for each seed for comparisons with the populations
studied by Galetti et al. (2013).
TESTING AND ANALYSIS
The comparisons between densities of palms in
fruiting, number of reproductive structures emitted
by reproductive palms, number of fruit formed
per infructescence, kilograms fruit produced per
hectare (ha), average seed size, bird richness,
wildlife actions (percentage of pulp, predation,
dispersion, etc.), and percentages of seed
germination and seedling mortality, were tested
using 95 % condence intervals (CIs; t distribution).
Comparisons between monthly frequencies of
species and wildlife families were tested by 95 %
CIs, which were obtained by resampling (1,000
bootstraps) using R software (R Development
Core Team 2015). The relationships between the
monthly frequencies of the wildlife families and
the percentages of fruit transport, pulping, and
intact fallen fruit were calculated using the Pearson
correlation, and statistical signicance (95 %) was
determined using the t test (t = r / √((1 - r2)/(n-2)).
RESULTS AND DISCUSSION
QUANTIFICATION OF FRUIT PRODUCTION
The study area had an average density of 231 ± 33
reproductive palms/ha; however, not all individuals
reproduced annually. In the 2008 fruiting season,
228 ± 33 palms/ha produced infructescences,
while in 2009, only 133 ± 36 palms/ha produced
infructescences. Considering the 11 recorded
fruiting seasons, the average number of palms
in reproduction per ha/year was 173 ± 37, with a
minimum value of 75 palms/ha/year in 2005 and
a maximum value of 266 palms/ha/year in 2015.
Annual variations in the density of reproductive
palms were also observed by Mantovani and
Morellato (2000), who registered variations
between 60 and 109 palms/ha.
Annual variations were observed in the number
of infructescences with ripe fruit per plant. In 2008,
considering only the reproductive palms, the average
number of ripe infructescences per plant was 1.47 ±
0.08, and in 2009, 0.86 ± 0.05 ripe infructescences
were registered per plant. These values were similar
to those observed by Silva Matos and Watkinson
(1998) and Mantovani and Morelatto (2000), with
variation between 0.63 – 1.8 infructescences/plant;
these authors did not distinguish between unripe
and ripe infructescences.
The number of ripe fruit produced per
infructescence also varied between years. In 2009,
despite a lower reproductive palm density, 2,789 ±
455 ripe fruit were registered per infructescence.
This number was higher than the observed value
for 2008 of 1,785 ± 220 fruit/infructescence.
These variations resulted in an annual production
that ranged from 599,310 fruit/ha/year in 2008
to 318,046 fruit/ha/year in 2009, or 1,031 and
547 kg/ha/year, respectively, (average 790 kg/
ha/year) when considering that each fruit weighs
approximately 1.72 g (Fleig and Rigo 1998).
The average number of fruit produced per
infructescence, as observed in other studies,
was quite variable (from 873 to 3,365 fruit/
infructescence; Silva Matos and Watkinson 1998,
Mantovani and Morellato 2000, Calvi and Piña-
Rodrigues 2005). This variation resulted mainly
from the different methodologies applied in the
estimates and makes comparisons with these

JULIANO Z. DA SILVA and MAURÍCIO S. DOS REIS CONSUMPTION OF Euterpe edulis FRUIT BY WILDLIFE
An Acad Bras Cienc (2019) 91(1) e20180537 6 | 20
studies of questionable value. However, based on
the number of reproducing plants in the 11 fruiting
seasons recorded in the present study, we estimated
production variation using the average number
of ripe infructescences emitted per reproductive
palm (1.17) and the average number of fruit
formed by infructescences (2,175) observed in
2008 and 2009. These estimates resulted in an
average production of 759 ± 162 kg/ha/year, with a
minimum production of 330 kg/ha/year registered
in 2005 and a maximum production of 1,163 kg/
ha/year registered in 2015. It should be noted
that the condence interval obtained (162 kg/ha/
year), represented 21.4 % of the average annual
production (equivalent to the production of 37
palms). This nding indicates that the wildlife is
naturally adapted to this fruiting oscillation.
IDENTIFICATION OF WILDLIFE AND
QUANTIFICATION OF THEIR ACTIVITIES
Fifty-nine vertebrate species that interacted with E.
edulis fruit and seeds (Campos et al. 2012, Galetti et
al. 2013) were recorded in Ibirama National Forest
(Table SI). However, only 31 species consumed E.
edulis fruit and seeds in the study area (Tables I, II
and SI). This number surpassed other studies, even
when considering the richness of birds alone (28
species; Silva Matos and Watkinson 1998, Galetti
et al. 1999, Fadini and Marco Jr 2004, Cerisola et
al. 2007, Fadini et al. 2009, Campos et al. 2012,
Omote et al. 2014, Cazassa et al. 2016, Silva et
al. 2017). The richness of frugivorous birds (Table
SI - total: 43, sporadic: 7, moderate: 6, extensive
frugivory: 6; regurgitate seeds: 19, defecate seeds:
3, seed predators: 7, and pulp consumers: 14) and
the average size of seeds regurgitated by these birds
(11.49 ± 0.9 mm) allowed us to classify the FLONA
de Ibirama as among the best areas considered as
non-defaunated (total richness: 36 ± 4.5, sporadic:
5.9 ± 0.9, moderate: 6.3 ± 0.5, extensive frugivory:
5.1 ± 1.1; regurgitate seeds: 17.3 ± 1.9, defecate
seeds: 1.5 ± 0.4, seed predators: 3.5 ± 0.9, and pulp
consumers: 13.7 ± 1.8; seed size: 11.39 ± 0.4 mm)
as described by Galetti et al. (2013).
In ripe infructescences, 6,956 interactions
were observed over 1,551 monitoring days. These
observations resulted in the identification of 24
species (Table I), mostly birds (23), results similar
to observations in other studies (Galetti et al. 1999,
2013, Silva et al. 2017). Monitoring of unripe
infructescences did not result in any interactions
despite including 9 palms and 126 monitoring
days. This nding indicated little interest among
vertebrates, a fact also suggested by the results of
installed collectors, as described below.
Of the total fruit used by vertebrates in
the infructescences, 30.4 % were pulped or
regurgitated, 62.6 % were dispersed, and 6.9 %
were knocked down during the feeding attempt
(Table I). The Turdidae family was most frequently
observed in infructescences (49.8 %; Table I), and
also represented the greatest fruit user (43.3 %),
observations similar to other studies (Silva Mattos
and Watkinson 1998, Galetti et al. 1999, Fadini et
al. 2009, Omote et al. 2014, Cazassa et al. 2016).
The main dispersers recorded were Turdus
flavipes, Selenidera maculirostris, and Turdus
albicollis. Together, these species accounted
for 87.6 % of primary dispersal (Table I). The
predominance of regurgitated seeds (97 %)
surpassed that of defecated seeds (2.1 %; by the
Cracidae family), and favors a higher percentage
of germination (Leite et al. 2012). Among the
dispersers, T. flavipes accounted for more than
one quarter of fruit consumed by vertebrates in the
infructescences and 41.7 % of primary dispersal
(Table I). This species was also reported as
among the most frequent visitors in other studies
(Galetti et al. 1999, Cerisola et al. 2007, Fadini
et al. 2009, Castro et al. 2012, Omote et al. 2014,
Cazassa et al. 2016). The importance of T. avipes
and its migratory behavior suggest that E. edulis
conservation needs to extend over a regional and
not just local population level.

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TABLE I
Wildlife recorded that fed on E. edulis infructescences, their frequencies of visitation, collected fruit per visit, percentage of fruit used, and probabilities and
percentages of primary dispersal (Disp.), dropping fruit (Dp.), predation (Pred.), and pulping, regurgitation, or defecation of seeds from infructescences (P, R or
D). These values represent the average for fruiting that occurred in 2013 and 2014.
Family Species Species Family Fruits/ %Fruits Used by Probability of: Percentages of:
Fq./MtzAFq./Mtz visit Species Family Disp. Dp. P/R/D Pred. Disp. Dp. P/R/D
Turdidae Turdus avipes 34.73 1.54 30.96 0.84 0.13 0.03 0 26.1 4.0 0.8
Turdus albicollis 14.15 1.42 11.63 0.85 0.11 0.04 0 9.9 1.2 0.5
Turdus amaurochalinus 0.20 1.16* 0.14 0.82 0 0.1 0.0 0.0
Turdus ruventris 0.60 1.59 0.56 0.85 0.15 0 0 0.5 0.1 0.0
Turdus leucomelas 0.10 49.78 0.97* 0.06 43.34 0.73 0 0.0 0.0 0.0
Ramphastidae Ramphastos dicolorus 1.81 3.92 4.11 0.92 0.07 0.01 0 3.8 0.3 0.1
Pteroglossus bailloni 0.10 3.60* 0.21 1 0 0 0 0.2 0.0 0.0
Selenidera maculirostris 10.91 12.82 3.24 20.47 24.79 0.92 0.06 0.02 0 18.8 1.2 0.5
Psittacidae Pyrrhura frontalis 15.37 1.57 13.92 0.03 0 0.97 0 0.4 0.0 13.5
Brotogeris tirica 0.76 1.21 0.53 0 0 1 0 0.0 0.0 0.5
Pionopsitta pileata 0.01 1.00 0.01 0 0 1 0 0.0 0.0 0.0
Triclaria malachitacea 0.42 16.56 1.42 0.34 14.81 0 0.06 0.94 0 0.0 0.0 0.3
Cotingidae Carpornis cucullata 1.16 1.38 0.93 0.92 0.07 0.02 0 0.9 0.1 0.0
Pyroderus scutatus 0.16 1.32 2.20* 0.20 1.13 1 0 0 0 0.2 0.0 0.0
Echimyidae Phyllomys sp. 19.02 19.02 1.30 14.31 14.31 0.02 0 0.98 0 0.2 0.0 14.1
Cracidae Penelope superciliaris 0.07 9.00 0.37 1 0 0 0 0.4 0.0 0.0
Penelope obscura 0.17 8.91* 0.89 1 0 0 0 0.9 0.0 0.0
Ortalis squamata 0.01 0.26 6.00 0.05 1.31 1 0 0 0 0.0 0.0 0.0
Tyrannidae Myiodynastes maculatus 0.01 0.01 2.00* 0.02 0.02 1 0 0 0 0.0 0.0 0.0
Trogonidae Trogon surrucura 0.01 0.01 1.00 0.01 0.01 1 0 0 0 0.0 0.0 0.0
Cardinalidae Habia rubica 0.04 0.04 2.50 0.06 0.06 0 0 1 0 0.0 0.0 0.1
Tityridae Tityra cayana 0.07 0.07 2.25* 0.09 0.09 0.89 0 0.1 0.0 0.0
Fringillidae Euphonia pectoralis 0.03 0.03 3.60* 0.06 0.06 0 0 1 0 0.0 0.0 0.1
Thraupidae Lanio melanops 0.06 0.06 2.00 0.07 0.07 0 0 1 0 0.0 0.0 0.1
Totals 62.6 6.9 30.4
A: Frequency per reproductive palm; * Values obtained by Galetti et al. (2013) because in our study, these species were very rare, allowing only a small sampling.

JULIANO Z. DA SILVA and MAURÍCIO S. DOS REIS CONSUMPTION OF Euterpe edulis FRUIT BY WILDLIFE
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TABLE II
Wildlife recorded on the soil surface that fed on E. edulis fruit and seeds, their frequencies of visitation, collected fruit or seeds per visit, percentage of fruit or
seeds used, and probabilities and percentages of secondary dispersal (Disp.), pulping, regurgitation, or defecation of seeds (P, R or D), buried seeds (Bur.), and
predation (Pred.). These values represent the average for fruiting that occurred in 2013 and 2014.
Family Species Species Family Fruits or % Used by Probability of: Percentages of:
Fq./MtzAFq./Mtz seeds/Visit Specie family Disp. P/R/D Bur. Pred. Disp. P/R/D Bur. Pred.
Turdidae Turdus albicollis 17.05 1.58 16.6 0.54 0.46 0 0 9.0 7.6 0.0 0.0
Turdus ruventris 3.69 20.74 2.15 4.9 21.5 0.82 0.18 0 0 4.0 0.9 0.0 0.0
Cotingidae Carpornis cucullata 0.46 0.46 1.00 0.3 0.3 1 0 0 0 0.3 0.0 0.0 0.0
Cricetidae Cricetidae 45.62 45.62 1.23 - 34.5 0.38 0.60 0 0.02 10.2*+3 20.7 0.0 0.6
Caviidae Dasyprocta azarae 15.90 15.90 2.34 22.9 22.9 0.92 0.07 0.01 0 21.1 1.7 0.2 0.0
Didelphidae Didelphis aurita 10.83 10.83 1.96 13.0 13.0 0.33 0.67 0 0 4.3 8.7 0.0 0.0
Momotidae Baryphthengus rucapillus 0.46 0.46 2.50 0.7 0.7 0.60 0.40 0 0 0.4 0.3 0.0 0.0
Formicariidae Chamaeza campanisona 3.46 3.46 1.93 4.1 4.1 0 1 0 0 0.0 4.1 0.0 0.0
Columbidae Geotrygon montana 0.92 0.92 2.00 1.1 1.1 0 0.33 0 0.67 0.0 0.4 0.0 0.8
Odontophoridae Odontophorus capueira 0.69 0.69 2.00 0.9 0.9 0 1 0 0 0.0 0.9 0.0 0.0
Cardinalidae Habia rubica 0.92 0.92 1.75 1.0 1.0 0 1 0 0 0.0 1.0 0.0 0.0
Totals 52.4 46.1 0.2 1.3
A: Frequency per reproductive palm; * Dispersal at close range < one meter from the fruits/seeds samples plot.

JULIANO Z. DA SILVA and MAURÍCIO S. DOS REIS CONSUMPTION OF Euterpe edulis FRUIT BY WILDLIFE
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The main fruit pulpers in infructescences
belonged to the Psittacidae family; they were
responsible for 47.1 % of pulped fruit. When
considering seed predators (Galetti et al. 2013), it
should be emphasized that they act only as pulp
consumers, i.e., species that predominantly remove
fruit pulp without carrying out the dispersal or
predation of the seeds, behavior already observed
by Laps R.R. (unpublished data).
Monitoring of vertebrate activities on E. edulis
fruit and seeds arranged at ground level totaled 434
records during 813 days of monitoring. Interactions
with at least 11 species were observed (Table II),
since rodents of Cricetidae were not identified.
These records characterized activities initiated
mainly on ripe fruit (430 records) and demonstrated
the relatively small interest of vertebrates in seeds.
Seeds were predominantly reserved for rodents
of the family Cricetidae and Geotrygon montana
(Columbidae) that predated 1.3 % of the amount of
fruit and seeds used by vertebrates at ground level.
Contrary to what was observed in
infructescences, mammals were the predominant
actors on the soil with respect to frequency of
interactions (72.4 %) and percentage of fruit used
(70.5 %). Here, we highlight the activities of
rodents of the Cricetidae and Caviidae families that
together used more than half of the available fruit
(Table II).
Vertebrate activities relative to fruit and seeds
arranged on the soil were predominantly involved
in secondary dispersal (52.4 %) and pulp (46.1 %;
Table II). However, dispersal was of a low quality,
generally related to the pulp at distances of less than
1 m from the samples (54.2 %). Greater distances
were only reached by seed-regurgitating birds
(T. albicollis, T. ruventris, Carpornis cucullata,
and Baryphthengus rucapillus) and by rodents
of the Cricetidae family, which presented a low
probability of seed predation (2 %; Table II).
USE OF FRUIT IN INFRUCTESCENCES
Through an analysis of fallen fruit in collectors
(Table III; Figure 1), we observed that an average
of 76.5 % of fruit were used directly from
infructescences (39.4 % transported, 25 % pulped,
0.5 % predated, 1.9 % insect drills, 6.9 % fungi,
2.8 % scratched or caught and lost). Contrary to
expectations, a very similar proportion of total
available fruit was utilized in 2008 and 2009, 77.9
± 3.4 % and 73.3 ± 9 %, respectively, even when
the amount of ripe fruit produced in 2009 was 47 %
lower than that observed in 2008.
Discounting the percentage destroyed by insects
and fungi (8.8 %; Table III), since they could not be
recorded by the camera traps, and correcting the
proportion of the fruit with the new total (67.7 %),
which expresses vertebrate activity, we calculated
the following proportions: transported: 58.2 %,
pulped: 36.9 %, predated: 0.7 %, and scratched:
4.1 %. These results were generally congruent with
those of the estimated percentages through the
installation of the camera traps (transported: 62.6
%, pulped: 30.4 %, predated: 0 %, and scratched:
6.9 %; Table I), and these ndings suggest small
variations between the fruiting seasons (collectors
considered the average of 2008 and 2009, while
camera traps considered the average of 2013 and
2014).
As shown in Table III, the percentage of fruit
with fungi and intact fruit tend to decrease, while the
percentage of transported fruit increased between
May and July. These results may be associated
with the onset of austral winter. During winter, the
region is drier than summer, which could contribute
to a reduction in fungal attacks. The changes in
percentages of intact and dispersed fruit (Table
III) were related to changes in the composition of
the wildlife (Figure 2a). Seed dispersers (mainly
Turdidae and Ramphastidae) predominated until
July, accounting for 84.7 % (CI: 78.3 - 90.6 %) of
the interactions (non-dispersers: 15.3 %, CI: 9.3 -

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TABLE III
Average percentage of wildlife and microorganismal activity on ripe E. edulis infructescences during the 2008 and 2009 fruiting seasons.
Period Pulped Scratched Predated
Vertebrates
Predated
Drilling
Dry and
cracked Intact Predated
Fungi Transported DestroyedaAble to
Germinateb
Used
wildlifec
May/2008 11.0 0.7 0.7 1.3 8.5 13.8 15.5 48.5 26.0 74.0 77.7
n=12 (3.9 – 18.0)d(0.2 – 1.2) (0.5 - 0.9) (0.3 - 2.3) (2.8 - 14.2) (12 - 15.7) (7.4 - 23.6) (41.9 - 55.1) (15.9 - 36) (63.9 - 84) (72.5 - 82.8)
June/2008 7.3 2.3 1.0 2.9 8.1 9.3 13.9 55.3 25.9 74.2 82.7
n=12 (2.9 – 11.7) (1.4 – 3.1) (0.2 - 1.7) (1.6 - 4.3) (4.6 - 11.6) (6.8 - 11.8) (5.7 - 22.1) (44.5 - 66.1) (14.5 - 37.4) (62.7 - 85.6) (77.6 - 87.7)
July/2008 8.3 2.4 1.7 3.5 9.4 9.0 10.4 55.5 25.0 75.2 81.8
n=13 (0.5 – 16.1) (1.5 – 3.2) (-0.3 - 3.6) (1.6 - 5.4) (5.4 - 13.3) (6.1 - 11.8) (6.3 - 14.4) (46.4 - 64.6) (16.6 - 33.2) (66.8 - 83.4) (76.1 - 87.3)
August/2008 21.4 2.4 0.3 1.7 2.3 14.0 6.0 51.9 10.3 89.7 83.7
n=13 (12.2 – 30.7) (1.8 – 3.0) (0.2 - 0.4) (0.3 - 3.2) (0.7 - 3.9) (4.3 - 23.6) (1.7 - 10.3) (36.1 - 67.6) (4.6 - 16.1) (83.9 - 95.4) (74.1 - 93.4)
September/2008 32.5 1.8 0.5 3.8 1.9 21.7 5.4 32.3 11.6 88.3 76.3
n=17 (22.9 – 42.1) (0.7 – 2.9) (-0.4 - 1.5) (-0.2 - 7.9) (0.6 - 3.3) (11.3 - 32.1) (-0.9 - 11.7) (21.1 - 43.4) (3.8 - 19.6) (80.4 - 96.2) (65.8 - 87)
October/2008 41.9 1.1 0.0 0.5 1.7 30.7 2.6 21.6 4.8 95.3 67.7
n=7 (28.1 – 55.6) (0.7 – 1.4) (0 - 0.1) (-0.3 - 0.1) (0.2 - 3.2) (16.4 - 45) (-1.7 - 7) (7.8 - 35.3) (-1.7 - 11.4) (88.6 - 101) (53.7 - 81.7)
November/2008 38.4 1.1 0.1 2.7 1.2 35.2 2.5 18.9 6.5 93.6 63.6
n=6 (18.9 – 58.0) (0.4 – 1.7) (-0.1 - 0.2) (-1.9 - 7.3) (0.6 - 1.8) (17 - 53.3) (-1.2 - 6.1) (8.3 - 29.5) (-1.2 - 14.1) (85.9 - 101) (45.5 - 81.8)
Mean 2008 21.0 1.8 0.7 2.5 5.0 17.1 8.6 43.2 16.9 83.1 77.9
n=80 (16.8 - 25.2) (1.5 - 2.1) (0.3 - 1.1) (1.6 - 3.5) (3.7 - 6.4) (13.7 - 20.6) (6.3 - 11) (38.3 - 48.1) (13.4 - 20.4) (86.6 - 79.6) (74.5 - 81.2)
July/2009 39.45 7.65 0.18 0.1 0.2 1.9 0.4 50.2 0.8 99.2 98.0
n=9 (27 - 52) (4 - 12) (0 - 0.4) (0 - 0.1) (0.1 - 0.2) (0.7 - 3.1) (-0.1 - 0.9) (36 - 65) (0.2 - 1.3) (99 - 100) (97 - 99)
October/2009 35.4 8.0 0.1 0.5 0.9 17.5 3.0 34.6 4.5 95.5 81.6
n=9 (19 - 52) (5 - 11) (0 - 0.2) (0 - 0.9) (0.4 - 1.4) (-2.3 - 37) (-0.1 - 6.2) (17 - 52) (1.1 - 7.9) (92 - 99) (61 - 102)
November/2009 29.4 5.2 0.1 1.0 0.5 42.6 1.9 19.4 3.5 96.5 56.9
n=6 (13 - 46) (2 - 8) (-0.1 - 0.3) (0 - 2) (0.1 - 0.9) (25 - 61) (0.1 - 3.6) (7 - 32) (1.7 - 5.2) (95 - 98) (39 - 75)
December/2009 31.6 1.1 0.1 0.5 1.1 45.4 5.2 15.2 6.8 93.2 53.5
n=10 (18 - 45) (0.6 - 1.5) (0 - 0.2) (0.2 - 0.7) (0.4 - 1.8) (32 - 58) (1.9 - 8.4) (6 - 24) (2.8 - 10.8) (89 - 97) (40.3 - 67)
Mean 2009 34.3 5.4 0.1 0.5 0.7 26.0 2.7 30.3 4.0 96.0 73.3
n=34 (28 - 41) (4 - 7) (0.1 - 0.2) (0.2 - 0.7) (0.4 - 0.9) (17 - 35) (1.5 - 4) (23 - 38) (2.5 - 5.5) (95 - 98) (64 - 82)
Mean Annual 25 2.8 0.5 1.9 3.7 19.8 6.9 39.4 13.1 86.9 76.5
n=114 (21.4 - 28.6) (2.3 - 3.4) (0.2 - 0.8) (1.2 - 2.6) (2.7 - 4.7) (16 - 23) (5.1 - 8.7) (35.1 - 43.6) (10.3 - 15.8) (84.2 - 89.7) (73 - 80)
a Destroyed: predated + dry and cracked; b Able to Germinate: pulped + scratched + transported + intact; c Used wildlife: pulped + scratched + predated + transported. n =
number of infructescences evaluated; d Condence intervals at 95 %.

JULIANO Z. DA SILVA and MAURÍCIO S. DOS REIS CONSUMPTION OF Euterpe edulis FRUIT BY WILDLIFE
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21.7 %). From August onwards, pulp consumers,
mainly Psittacidae and Echimyidae, predominated
and reached 62.9 % (CI: 52.5 - 72.1 %) of the
interactions (dispersers: 37.1 %, CI: 27.8 - 47.3
%). This wildlife shift resulted in changes in the
monthly percentages of transported, pulped,
and fallen intact fruit. Overall, there were high
correlations between the Turdidae family and the
percentages of fruit transported (r = 0.85) or fallen
intact (r = -0.87), and between the Psittacidae and
Echimyidae families and the percentage of pulped
fruit on the infructescences (r = 0.88 for both).
The largest change in wildlife composition
occurred in the Turdidae family (Figure 2a),
mainly represented by T. albicollis and T. avipes,
both of which reduced their frequencies after
August (Figure 2b). Part of this reduction could be
explained by the gradual migration of T. avipes,
which occurs during late winter and spring, when
it moves from the southern mountainous regions to
the north to the Espirito Santo State (Sick 1997), in
addition to migrating along the altitudinal gradient
following the fruiting of E. edulis (Castro et al.
2012). However, T. albicollis is a resident species,
a fact that indicates that it naturally uses other
resources after August, as corroborated by Castro
et al. (2012), who observed that the fruiting of E.
edulis inuences the abundance and migration of
T. avipes, but not T. albicollis, between dierent
altitudes and forest typologies. These data reect
concerns about fruit management between April
and July in Southern Brazil, in particular with
respect to the Turdidae family, its feeding habits,
and population dynamics, owing to annual
variations in fruit production.
Figure 1 - A schematic overview of the wildlife and microorganismal activities that occurred during E. edulis fruiting (from 114
infructescences and 1,253 samples of unripe, ripe, dried, and pulped fruit placed on the ground, with 73,129 fruit per seed).

JULIANO Z. DA SILVA and MAURÍCIO S. DOS REIS CONSUMPTION OF Euterpe edulis FRUIT BY WILDLIFE
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Of the total amount of fruit, only 19.8 % fell
to the ground intact (Table III) with no evidence
of animal activity, and these fruit were also free of
fungi and insect drills. This result highlights the
importance of this food source for wildlife, mostly
birds, that feed directly on infructescences (Table
I), as discussed by Galetti et al. (2000), Reis and
Kageyama (2000), and Galetti et al. (2013). It is
also evident that the supply of pulp on the ground
is proportionately low, corresponding to only 22.6
% of what is produced in an infructescence (19.8
% intact + 2.8 % scratched) or only 179 kg/ha/
year, considering the average production of 2008
and 2009 (790 kg/ha/year). Therefore, the fruit is
characterized as a scarcer resource on the ground
and concentrated beneath reproductive palms.
For natural regeneration of the species, 86.9 %
of the total amount of fruit produced remained and
were categorized into fruit viable for germination
{25 % pulped, 19.8 % intact, 2.8 % scratched, and
39.4 % transported (dispersers are regurgitators
and seed defecators); Figure 1}. This represents an
average of 399,050 seeds/ha.
Figure 2 - Monthly average frequency of the: a) families and b) species recorded that fed on E.
edulis infructescences during April to July and August to November during the 2013 and 2014
fruiting seasons. Condence intervals (95 %) are indicated.

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USE OF FRUIT ON THE SOIL
For wildlife that feed on the ground, dierent types
of fruit are available {unripe (green), ripe, pulped,
and dry pulp}. Only 8 % of the ripe fruit was
unused (Table IV; Figure 1). In contrast, unripe,
pulped, and dried fruit were barely utilized by
wildlife. When analyzing the percentage of pulped
and transported fruit, as shown in Table IV, fruit
preference was evident, since 98 % of pulping
and 86 % of dispersing that occurred in the soil
originated from ripe fruit pulp.
The behavior of wildlife on pulped fruit was
very similar to that observed for unripe and dry
fruit, where 96.3 % of the fruit showed no animal
activity (Table IV). However, pulped and unripe
fruit contributed 13 % to secondary dispersal, and
thus represented, 5.1 and 8.2 %, respectively, of
dispersed seeds. As the main food resource oered
by pulped fruit is the seed’s endosperm, the seeds
would be destroyed and thus contribute very little
to secondary dispersal. This supposition was
corroborated by records obtained with the camera
traps, which only recorded activities on seeds
that resulted in predation. In addition, unripe fruit
reaches the ground as immature fruit with insect
drills or fungi in the endosperm.
Based on the information presented in Table
IV, the importance of ripe fruit for wildlife that
feeds on the ground and for secondary dispersal is
evident. Furthermore, the results obtained provide
an understanding of the sensitivity of this system
since the amount of ripe fruit that reached the forest
floor was a relatively small proportion (22.6 %
produced by an infructescence), and of these fruit
only 8 % were not consumed. That is, only 1.8 %
of ripe fruit produced were not used by animals
(14.2 kg/ha/year or the equivalent of 3.8 ripe
infructescences/ha/year).
Secondary dispersal involved the transport
of 10.9 % of the total produced fruit (Figure 1),
and originated from the actions of wildlife on soil
fruit with ripe pulp (22.6 %: 19.8 % intact and 2.8
% scratched; Table III; Figure 1). The 10.9 % of
secondary dispersal is probably underestimated
because the wildlife that feed on the ground can
still move the fruit and seeds via primary dispersal.
However, this percentage will not be much greater,
since primary dispersal animals carry the fruit but
only because of their interest in the pulp. The fruit
are then likely to be discarded after being pulped.
As shown in Table IV, few seeds left on the soil
were dispersed by wildlife.
CONSUMPTION OF FRUIT AND PERCENTAGE OF
SEEDS AVAILABLE FOR NATURAL REGENERATION
Seeds available for natural regeneration originated
from 19.8 % intact fruit, 25 % pulped fruit, 2.8
% scratched fruit, and 39.4 % dispersed fruit (a
total of 87 %; Table III; Figure 1). However, these
percentages are reduced due to ecological processes
that occur on the soil, where seeds are subjected to
pulping, predation, transport, and rotting caused by
microorganisms. These reductions are outlined in
Figure 1 and detailed in the Appendix.
Including all losses and reductions, 70.4 % (19
% + 21.5 % + 29.9 %) of seeds could germinate. Of
the percentage destroyed, 13 % occurred by direct
wildlife activity on infructescences (Table III;
Figure 1). The destruction of the remaining 16.6
% of seeds occurred mainly in the soil and during
dispersal (13.3 % with fungal attacks and 3.3 % due
to predation and rot). These results are summarized
in Figure 1, which shows 70.4 % of seeds were
capable of germination and 29.6 % were destroyed
(20.2 % by fungus, 1.9 % by insect larvae, 3.8 %
by predation and microorganisms, and 3.7 % as a
result of dry and cracked fruit).
The estimated percentage of seeds that could
germinate (70.4 %) was very close to that observed
for seeds evaluated in the greenhouse (78.8 ± 5.1
%). The percentage of destroyed seeds (21.2 ±
5.1 %) was possibly related to fungal attack, since
the same percentage was observed by the internal

JULIANO Z. DA SILVA and MAURÍCIO S. DOS REIS CONSUMPTION OF Euterpe edulis FRUIT BY WILDLIFE
An Acad Bras Cienc (2019) 91(1) e20180537 14 | 20
TABLE IV
Average percentage of wildlife and microorganismal activity on the dierent classes of available fruit on the soil during
the 2008 and 2009 fruiting seasons.
Wildlife and/or microorganism actions
Class of Intact Pulped Predated Pred. dril. Transported Rotten
Fruit %CFa%CAb%CF %CA %CF %CA %CF %CF %CA %CF %CA
Dry pulpc83.3 29.7 0.4 1.0 0.4 17.4 0 0.3 0.5 15.6 92.3
CI(±) 12,9 0.5 0.5 0 0.2 12.7
Green fruitsd93.0 33.1 0.6 1.5 0.9 39.1 0.2 4.8 8.2 0.5 3.0
CI(±) 11.0 1.3 1.6 0.5 10.0 1.3
Ripe fruitse8.0 2.9 40.2 97.6 0.5 21.7 0 50.7 86.2 0.6 3.6
CI(±) 4.3 9.1 0.5 12.2 0.8
Pulped fruitsf96.3 34.3 x x 0.5 21.7 0 3.0 5.1 0.2 1.2
CI(±) 3.8 0.5 3.7 0.3
Mean 70.2 14.0 0.6 0.1 14.7 4.2
Pred. dril.: predated by drilling insects, a %CF: Percentage within the class of fruit, b %CA: Percentage within class of wildlife
activities (%CA is a proportion of the total %CF for all classes of fruits). c Dry pulp: 15 evaluations involving 346 samples and
9,269 fruits. d Green fruits (unripe): 8 evaluations involving 47 samples and 2,551 fruits. e Ripe fruits: 17 evaluations involving
251 samples and 11,244 fruits. f Pulped fruits: 18 evaluations involving 609 samples and 50,065 fruits. CI(±): Condence intervals
at 95 %.
analysis of ripe fruit that fell in the collectors
(Appendix). The dierences between germination
percentage (70.4 and 78.8 %) are likely related
to the environmental conditions observed in the
greenhouse (no soil level predation) as well as the
characteristics of the seeds used (ripe fruit pulped
by wildlife) that exclude attacks of insect drills
observed in unripe fruit and cracked and dry fruit
recorded in infructescences.
According to our results, the largest bottleneck
related to the regeneration of E. edulis is not
related to seed destruction before germination,
since 322,909 seeds/ha (70.4 %) could germinate,
a much higher percentage than the number of
seedlings found in class I in 2009 in the study area
(90,876 ± 31,290 seedlings/ha). Instead, the highest
mortality occurred after seed germination during
seedling establishment, where between August
2008 and February 2009 (7 months), 63.2 ± 7 %
of the germinated seeds (seedlings) on the samples
arranged on the soil had already died. Other studies
corroborated our results and reported large class
I mortalities (between 47.2 % and 55.1 %) in
their rst year of life (Conte et al. 2000, Reis and
Kageyama 2000, Silva J.Z., unpublished data).
Considering the dynamics of E. edulis natural
regeneration, where seedling density is greater
closer to reproductive palms, and the limiting
effects imposed by density (Silva Matos and
Watkinson 1998, Reis and Kageyama 2000), it
is likely that only dispersed seeds will be able to
replace the current reproductive palms. Of the 70.4
% seeds that may germinate, these results imply
that only 40.8 % are dispersed to conditions that
favor establishment (29.9 % from primary dispersal
and 10.9 % from secondary dispersal; Figure 1).
The remaining 29.6 % are not dispersed and will be
subject to new fruiting seasons and seed production
from the reproductive palms as well as the limiting
conditions imposed by the location itself, such
as falling leaves, spathes, and bunches. Studying
the dynamics of natural regeneration, Tonetti and
Negrelle (2001) observed that the fall of leaves or
branches is responsible for annual mortality of 20.6

JULIANO Z. DA SILVA and MAURÍCIO S. DOS REIS CONSUMPTION OF Euterpe edulis FRUIT BY WILDLIFE
An Acad Bras Cienc (2019) 91(1) e20180537 15 | 20
% of E. edulis seedlings. In this sense, we emphasize
that local conditions may have a predominant eect
on seed quantity that will remain in management
systems.
IMPLICATIONS FOR MANAGEMENT AND
CONSERVATION
Our results showed that E. edulis fruit are an
important resource for wildlife, which use most
of the available resources irrespective of whether
production during the year is high or low. These
findings make it possible to conclude that
sustainable management in natural populations is
not possible by simply identifying surpluses, since
they are very small. On the other hand, the large
variation in the number of fruit produced annually
(47 %) leads us to question how wildlife would
react to these variations, or even their management,
since a) E. edulis may only be a preferred
resource and there are other available resources
or b) whether there are periods of limited wildlife
development due to lack of food caused by low
fruitings or even fruit harvesting. To answer these
questions, additional studies are needed, mainly
aimed at identifying variations caused by years of
high and low E. edulis fruit production such as: a)
population uctuations of wildlife, both in size and
composition, resulting from oscillating food supply;
b) changes in the percentages consumed annually
by dierent species, since such percentages could
be a reflection of competition for fruit (Turdus
spp. were quite aggressive among themselves) and
actions could be directed to species more aected
by lack of fruit removal; c) characterize the fruiting
phenology of other species in order to demonstrate
the importance of E. edulis fruit throughout the
year, and other possibilities of food for wildlife.
Contrary to our hypothesis, we could not
conrm that the mere identication of E. edulis
fruit surpluses would be sufficient to initiate a
management program for fruit; however, we can
report several observations that might suggest a
direction for use and conservation strategies:
a) The volume of ripe fruit produced per
hectare can vary considerably between years;
in addition, the observed variations were
recurrent, and this phenomenon suggests that
wildlife is accustomed to some intensity of this
oscillation. Thus, harvesting fruit only in years
where the production would surpass the annual
average may cause minimal impact. The
harvest should be limited to the amount that
exceeds the annual average. Considering the
application of this method on the production
estimates (11 fruitings), it would be possible
to manage an average of 94 ± 90 kg/ha/year.
Based on the prices paid to the extractor for
E. edulis fruit per kilogram in 2017 in Santa
Catarina State (R$ 3.50/kg; Companhia
Nacional de Abastecimento 2017), it would be
possible to obtain an average of R$ 329/ha/
year. Although this is a conservative estimate,
it represents R$ 3,290/year on a small property
(10 ha) or R$ 152,327/year in a big area such
as the FLONA (463 ha).
b) The quantity of ripe fruits produced/ha/year is
a key point in determining the balance between
harvesting, feeding of wildlife, dispersal and
natural regeneration. Unripe or dry pulp fruits
are little used as food by the fauna, and their
seeds have reduced germination, contributing
little to natural regeneration. In addition, seeds
(pulped fruit) are poorly dispersed, limiting
the contribution of secondary dispersal.
c) Although many vertebrates are associated with
E. edulis (Galetti et al. 2013), the presence
of these species does not guarantee they will
interact with the fruit. In our study, only 52.5
% of species interacted with E. edulis and
used its fruit and seeds as food. Thus, we
believe that oristic composition and faunal
interactions also have an important eect on

JULIANO Z. DA SILVA and MAURÍCIO S. DOS REIS CONSUMPTION OF Euterpe edulis FRUIT BY WILDLIFE
An Acad Bras Cienc (2019) 91(1) e20180537 16 | 20
the identication of species that depend on E.
edulis fruiting.
d) Most pulp is consumed from infructescences
(76.5 %), and this fact dictates the direction
of future studies, since many currently only
include monitoring of activities that occur
on the soil that represent the minority of
interactions.
e) The most active species in the consumption and
dispersal of seeds (T. avipes) is a migratory
bird, and this fact suggests the need for a greater
reach for E. edulis conservation strategies.
The protection of an E. edulis population also
depends on the conservation status of other
populations for which T. avipes migrates or
moves along the altitudinal gradient.
f) The composition of wildlife changes during
the fruiting months, and this fact indicates
that the importance of E. edulis fruit can be
quite variable, depending on the needs of
each species throughout the fruiting period. In
addition, the change in wildlife composition
changes dispersal percentages. Therefore, after
August, it is likely that fruit management could
benet the natural regeneration of E. edulis
because it would reduce the density of seedlings
and seeds in the vicinity of the reproductive
plants. In this period, dispersal decreases and
larger amounts of seeds accumulate around
the palms; this phenomenon tends to lead to
higher seedling mortality (Silva Matos and
Watkinson 1998, Silva Matos et al. 1999, Pizo
and Simão 2001).
g) The percentage of dispersed seeds, discounting
the destroyed seeds, totaled 40.8 %, 29.9 % of
which resulted from primary dispersal and 10.9
% from secondary dispersal. It is important
to note that most (97 %) of the dispersed
seeds were regurgitated, which favors higher
germination percentages, as discussed by Leite
et al. (2012). Secondary dispersal is of low
quality and is generally related to pulping at
distances of less than 1 m from fruiting palms
(54.2 %) and performed by small rodents
(Cricetidae) and by Dasyprocta azarae.
h) The estimate of the percentage of seeds that
could germinate in the forest was high (70.4
%), and seedling mortality in its rst year of life
reached 63.2 % in only 7 months. In this sense,
local conditions in which natural regeneration
is found may be of greater importance to
the number of growing seedlings rather than
the quantity of seeds that will remain in
management systems.
ACKNOWLEDGMENTS
We would like to thank the team of environmental
analysts at the National Forest Ibirama: Flavio
Zanchetti, Homero de Oliveira Salazar Filho and
Marcela Xavier Machado (Chico Mendes Institute
for Biodiversity Conservation – ICMBio). This
study was supported by the Conselho Nacional
de Desenvolvimento Científico e Tecnológico
(CNPq), 309128/2014-5 to Maurício S. Reis,
Coordenação de Aperfeiçoamento de Pessoal de
Nível Superior (CAPES), postdoctoral fellowship
given to Juliano Z. da Silva, and the Fundação de
Amparo à Pesquisa e Inovação do Estado de Santa
Catarina (FAPESC) - PRONEX/2780/2012-4.
AUTHOR CONTRIBUTIONS
J.Z.S. and M.S.R. contributed to the design of
the research. J.Z.S. performed the measurements,
processed the experimental data, drafted the
manuscript and designed/produced the figures.
M.S.R was involved in supervised the Work. All
authors discussed the results and commented on
the manuscript.
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SUPPLEMENTARY MATERIAL
Table SI - Wildlife recorded in Ibirama National Forest
(Management PlanA and this study) associated with the
consumption of Euterpe edulis fruit and pulp of by Galetti et
al. (2013), Campos et al. (2012), and this study.
Figure S1 – a) Model of fruit collectors installed in the study
area. b) and c) Using camera traps to monitor infructescences.
d) Samples that contained fruit or seeds arranged on the soil.
Drawing of Euterpe edulis adapted from Henderson (2000).
Figure S2 - Wildlife recorded interacting with Euterpe edulis
fruit and seeds in Ibirama National Forest. * Image by Homero
O. Salazar; other images by Juliano Z. Silva.
APPENDIX
DETAILS OF THE CONSUMPTION OF FRUIT AND
THE PERCENTAGE OF SEED AVAILABLE FOR
NATURAL REGENERATION FROM WILDLIFE
ACTIVITIES OCCURRING AT GROUND LEVEL.
Seeds available for the natural regeneration process
basically have as origin the following: 19.8 % intact
fruit, 25 % pulped fruits, 2.8 % scratched fruit and
39.4 % dispersed fruits = 87% (Table III; Figure
1). However, these percentages suer even greater
reductions owing to the ecological processes
occurring on the soil where seeds are subjected to
pulping, predation, transport and rotting caused by
microorganisms.
Of the rst 25 % of fruit reaching the ground
as pulped, 96.3 % will remain intact, 3 % will
be transported and 0.7 % will be destroyed by
predation and rot (Table IV; Figure 1). However,
for 3 % of seeds that are transported, the motivation
is the endosperm, since it is pulped. Therefore,
the chance of predation is high. For this reason,
these seeds are considered destroyed, reducing the
percentage from 25 to 24.1 %. However, these fruits
are also subject to fungal contamination, and unlike
the value obtained for intact fruits, this value has
yet to be corrected. To do this, we used the average
percentage of fruits with fungal attacks found in
each collector during the period in which the ripe
fruit was subjected to transport and pulping, where
the value found was 21 %. This correction resulted
in 19 % participation in the regeneration process
(Figure 1).
Ripe fruit reaching the ground was composed
of 19.8 % fallen intact fruit and 2.8 % scratched
fruit (19.8+2.8 = 22.6 %). From Table IV and
Figure 1, it can be seen that 40.2 % will be pulped,
8 % will remain intact, 50.7 % will be transported
and 1.1 % will be lost through decay and predation.
The 40.2 % that will be pulped on the soil begins to
behave like pulped fruits that reach the ground, i.e.,
96.3 % contribute to regeneration and 3.7 % will be
destroyed. Thus, of the 40.2 % pulped fruits, only
38.7 % of the 22.6 % of ripe fruits that reach the
forest oor contribute to the regeneration process
(8.75 %). The 50.7 % of seeds transported from
the ground that contain pulp, as suggested by the
data in Table IV, will be pulped with preference.
Nevertheless, as shown by the data in Table IV, for
ripe fruit, 0.5 % will be predated, 0.6 % will rot,
and the remaining 98.9 % will be pulped (11.33
%). However, after being pulped, they will assume
the behavior observed for pulped fruit on the soil,
where, again, 96.3 % will contribute to regeneration
(10.91 %) and 3.7 % will be destroyed (0.42 %).

JULIANO Z. DA SILVA and MAURÍCIO S. DOS REIS CONSUMPTION OF Euterpe edulis FRUIT BY WILDLIFE
An Acad Bras Cienc (2019) 91(1) e20180537 20 | 20
In this regard, while 50.7 % of fruit will be
transported from the ground, only 48.8 % will
contribute to the regeneration process. Thus, of the
22.6 % of ripe fruits that arrive intact on the forest
oor, 95.5 % of the seeds (8 + 38.7 + 48.8) will
contribute to the regeneration process, i.e., 21.5 %
(1.8 + 8.8 + 10.9 – Figure 1).
Given that the main attraction associated
with primary dispersal is the pulp, 39.4 % of the
transported fruit in primary dispersal is likely to be
pulped. However, 0.5 % predation was observed
within collectors, which may also occur with
transported fruit. Thus, the percentage of fruits
transported that can germinate will be reduced to
39.2 %. These fruits will also be subject to fungal
attack, requiring a correction in value. In order to
do this, the average percentage of fruit infected with
fungus was found for each sample in each collector
during the period in which ripe fruit was subjected
to transport. The value was 21 %, which reduces
the value of 39.2 to 31 %. As they reach the ground,
pulped fruit will still suer from 0.5 % predation,
0.2 % from decay and 3 % from transport, which
is motivated by the endosperm of the seed, most
likely destroying them. Computing these losses to
3.7 %, the percentage of transported fruits able to
germinate will be 29.9 % (Figure 1).
Including all losses/reductions, 70.4 %
(19+21.5+29.9) represents total fruit destined
to become germinating seeds (Figure 1). The
destruction of 16.6 % occurred mainly on the
soil and during dispersal, and of this total, 13.3
% results from fungal attacks and 3.3 % from
predation and rot. These results are summarized
in Figure 1, which shows 70.4 % of seeds capable
of germination and 29.6 % of seeds destroyed
by fungal attach (20.2 %), insect larvae (1.9 %),
predation and microorganisms (3.8 %) and dry and
cracked fruits (3.7 %).