Accepted Manuscript

Scavenging as a pathway for plastic ingestion by marine animals

Ryan Andrades, Roberta Aguiar dos Santos, Agnaldo Silva Martins, Davi Teles,
Robson Guimarães dos Santos

PII: S0269-7491(18)33344-X
DOI: https://doi.org/10.1016/j.envpol.2019.02.010
Reference: ENPO 12179

To appear in: Environmental Pollution

Received Date: 21 July 2018

Revised Date: 28 January 2019
Accepted Date: 3 February 2019

Please cite this article as: Andrades, R., dos Santos, R.A., Martins, A.S., Teles, D., dos Santos,
Robson.Guimarã., Scavenging as a pathway for plastic ingestion by marine animals, Environmental
Pollution (2019), doi: https://doi.org/10.1016/j.envpol.2019.02.010.

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1 Scavenging as a pathway for plastic ingestion by marine animals

2 Running page head: Scavenging as a pathway for plastic ingestion

4 Ryan Andrades1, Roberta Aguiar dos Santos2, Agnaldo Silva Martins3, Davi Teles4, Robson
5 Guimarães dos Santos4,*

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7 Laboratório de Ictiologia, Departamento de Oceanografia e Ecologia, Universidade Federal
8 do Espírito Santo, Av. Fernando Ferrari, 514, Goiabeiras, Vitória, Espírito Santo, Brazil,

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9 29075-910;

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10 Instituto Chico Mendes de Conservação da Biodiversidade, Centro de Pesquisa e Gestão dos

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11 Recursos Pesqueiros do Litoral Sudeste e Sul, Itajaí - SC, Brazil;
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12 Laboratório de Nectologia, Departamento de Oceanografia e Ecologia, Universidade
13 Federal do Espírito Santo, Brazil;
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14 Instituto de Ciências Biológicas e da Saúde, Universidade Federal de Alagoas, Maceió -
15 AL, Brazil.
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17 Corresponding author:
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18 E-mail: robsongsantos@gmail.com

19 Tel/Fax: +55-82-32141651
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20 ABSTRACT

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22 Plastic pollution is prevalent worldwide and affects marine wildlife from urbanized beaches
23 to pristine oceanic islands. However, the ecological basis and mechanisms that result in
24 marine animal ingestion of plastic debris are still relatively unknown, despite recent
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advances. We investigated the relationship between scavenging behavior and plastic
26 ingestion using green turtles, Chelonia mydas, as a model. Diet analysis of C. mydas showed
27 that sea turtles engaging in scavenging behavior ingested significantly more plastic debris

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28 than individuals that did not engage in this foraging strategy. We argue that opportunistic
29 scavenging behavior, an adaptive behavior in most marine ecosystems, may now pose a

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30 threat to a variety of marine animals due to the current widespread plastic pollution found in
31 oceans.

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Keywords: Chelonia mydas; marine pollution; plastic pollution; marine debris; scavenging
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33 Scavenging behavior in sea turtles led to high rates and loads of plastic ingestion and may be an
important pathway for plastic debris ingestion by marine animals.
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35 INTRODUCTION

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37 Plastic pollution in the world’s oceans is ubiquitous (Barnes et al., 2009; Cózar et al., 2014;
38 Thompson et al., 2009), and its impacts on marine animals have been reported in multiple
39 studies (Browne et al., 2015; Gall and Thompson, 2015; Laist, 1997). Since the first attempt
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to produce a comprehensive list of species with plastic ingestion records made by Laist
41 (1997), the number of studies reporting plastic ingestion has increased dramatically (Gall and
42 Thompson, 2015). Despite the number of studies reporting plastic ingestion by marine

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43 animals, we still have gaps in our knowledge related to the impacts and causes of plastic
44 ingestion (Browne et al., 2015; Rochman et al., 2016; Lynch 2018). Plastic ingestion has

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45 been reported for a great variety of species that rely on different sensory organs to perceive
46 the environment and that have different foraging strategies, from zooplankton (Desforges et

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47 al., 2015) to whales (Stephanis et al., 2013); therefore, understanding the factors associated
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48 with plastic ingestion is a complex and challenging task. Despite the complexity of this topic
49 and the lack of apparent patterns, some studies have identified possible drivers behind plastic
50 consumption, such as visual cues (Ryan, 1987; Santos et al., 2016; Schuyler et al., 2014,
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51 2012), chemical cues (Allen et al., 2017; Savoca et al., 2016) and foraging strategies (Ryan,
52 1987; Santos et al., 2015a; Schuyler et al., 2012).
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53 In most marine ecosystems, food resources may be scarce and unpredictable in time
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54 and space. In such environments, opportunistic scavenging may be an adaptive behavior

55 (MacArthur and Pianka, 1966) regardless of the species diet and foraging strategy.
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56 Additionally, the large home ranges common in many marine animals may enhance the
57 chances that they will encounter scavenging opportunities (Kane et al., 2017). Indeed, the
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58 estimated prevalence of scavenging in marine food webs is higher than that in estuarine,
59 freshwater and terrestrial food webs (Wilson and Wolkovich, 2011). Facultative scavenging
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60 may act as an important stabilizing force in many food webs by making the food webs more
61 reticulated (McCann et al., 1998; Wilson and Wolkovich, 2011). Despite the importance of
62 facultative scavenging to the understanding of food webs, few studies were able to address
63 this issue, and it has been suggested that the prevalence of scavenging in food webs is
64 underestimated by 16-fold (Wilson and Wolkovich, 2011).

65 Opportunistic scavenging implies that foragers will be more prone to ingest a broader
66 range of prey types, and we may expect that foragers adopting this behavior may enhance

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67 their chances of mistaking plastic debris for food items. The relationship between broader
68 diets and plastic ingestion has been suggested for seabirds (Ryan, 1987), which may also
69 contribute to trophic transfer of plastics from prey to generalist predators (Farrel and Nelson,
70 2013; Lusher et al., 2015). Therefore, due to the high prevalence of facultative scavenging
71 expected to occur in marine animals, mainly in coastal systems and sea floors where carrion
72 availability is most common (Britton and Morton, 1994, Beasley et al., 2012, Kane et al.,

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73 2017), opportunistic scavenging may play a role in understanding the high numbers of marine
74 species ingesting plastic debris.

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75 In this study, we investigate the relationship between scavenging and plastic ingestion
76 using the green turtle, Chelonia mydas, as a model. Although green turtles are known to be

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77 the only species of sea turtles that have specialized in an herbivorous diet, a growing number
78 of studies have demonstrated foraging behavioral plasticity, with several records of animal

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79 matter ingestion (Cardona et al., 2009; González Carman et al., 2013; Reisser et al., 2013;
80 Santos et al., 2015a). Behavioral plasticity is a common attribute of organisms and plays a
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81 central role in optimal foraging theory (Ghalambor et al., 2010). The ingestion of animal
82 matter by sea turtles may be accidental (e.g., isopods; Santos et al., 2015b) or intentional
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83 through direct predation (e.g., jellyfish; González Carman et al., 2013) or scavenging (e.g.,
84 scavenging; Limpus et al., 2008).
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85 To assess the possible relationship between scavenging behavior and plastic ingestion,
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86 we evaluated the possible scavenging behavior of green turtles related to the ingestion of
87 cephalopods and the amount of ingested plastic. For this, we addressed the following
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88 questions and built a framework leading to evidence that scavenging behavior can promote
89 and enhance debris ingestion chances.
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91 1. Is squid ingestion by sea turtles a result of scavenging behavior?

92 2. Is scavenging behavior correlated with plastic ingestion?

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94 MATERIALS AND METHODS

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96 From 2009 to 2013, 295 dead, stranded green turtles were collected along the
97 Brazilian coast, from the states of Alagoas (09˚39’S, 35˚44’W) to Santa Catarina (27˚26’S,
98 48˚19’W) (Table S1). Analyses concerning feeding habits and debris ingestion were
99 performed, examining entire gastrointestinal systems to retrieve debris and cephalopod
100 remains and beaks. We quantified marine debris by weight (dry mass; 0.01 g digital scale)
101 and by number of items (only fragments longer than 0.5 cm were considered an item). We

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102 considered that debris smaller than 0.5 cm may be generated by the fragmentation of larger
103 items inside the turtle. Therefore, these items were accounted for only in the total weight, not

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104 in the total number of ingested items. However, plastic pellets (raw plastic material of plastic
105 products) were considered as an individual item, independent of size. All cephalopod beaks

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106 were identified based on beak morphology and comparisons to vouchers from the scientific
107 cephalopod collection of Centro Nacional de Pesquisa e Conservação da Biodiversidade
108 Marinha do Sudeste e Sul – CEPSUL, Brazil. Whenever possible (i.e., in unbroken beaks),
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estimations of mantle lengths and body mass of squid were made by performing linear
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110 regressions from measures of upper rostral length and lower rostral length of beaks following
111 specialized literature (Clarke, 1986; Lu and Ickeringill, 2002). Detailed information on diet
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112 and debris ingestion can be found in Santos et al. (2015a,b).

113 The necropsies and size measurements (curved carapace length – CCL) were
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114 performed by trained veterinarians using standard techniques (Wyneken, 2001). The necropsy
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115 started by removing the plastron. The major groups of pectoral muscles were evaluated and
116 removed, as well as the heart and liver. A midventral cut in the neck was made to expose the
117 esophagus, and the gastrointestinal tract was collected (from esophagus to rectum). Debris
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118 ingestion was assigned as the cause of death only when the turtles were underweight or
119 emaciated, the gastrointestinal tract (GIT) was blocked by debris, and when there was no
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120 evidence of another cause of death (for more details, see Santos et al. (2015a).
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121 We performed a Mann-Whitney test to evaluate whether sea turtles with and without
122 squid remains in their gastrointestinal tract differed in size (CCL). To test whether
123 scavenging behavior, inferred by squid ingestion, had any influence on debris ingestion
124 (number of items and weight), we performed a Mann-Whitney paired test comparing animals
125 with and without squid remains in their gastrointestinal tract.

126 We recognize that the sea turtles’ size (i.e., CCL) and foraging areas may have an
127 important role in debris ingestion. The size of a sea turtle may influence the amount of plastic

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128 that can be ingested by each individual (Wilcox et al., 2018), and the availability of plastic in
129 the environment and sea turtles’ foraging strategies (benthic or water column foraging
130 strategies; Santos et al. (2015b)) may vary among sea turtle foraging areas. Therefore, we
131 modeled the effect of foraging area, curved carapace length and scavenging behavior with
132 plastic ingestion using a hurdle regression model considering zero-inflated count data (Zeileis
133 et al., 2008). We choose this analysis due to the relative number of turtles that ingested squid

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134 in relation to those that did not. We modeled the effects of the abovementioned variables
135 regarding the presence of plastic debris and the amount of plastic debris ingested by each

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136 turtle (i.e., the number of plastic items). We tested three different probability distributions
137 (Poisson, negative binomial and geometric) for the models and selected the most plausible

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138 model according to the ∆AICc (Burnham et al., 2011). The analyses were performed with the
139 statistical software R (R Core Team, 2015) using the packages ‘pscl’ (Jackman, 2017) to run
140 the hurdle models and ‘MuMIn’ (Barton, 2009) to compare and select the best model (for the
141 analysis script, see Supplementary Material).
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143 RESULTS

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145 All analyzed turtles were considered neritic (i.e., coastal zone extending seaward to
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146 outer shelf edge depths) juveniles based on their size (mean CCL: 37.9; SD: 7.0; range: 26.1–
147 78.4 cm) and their diet items, composed mainly of benthic macrophytes (for detailed
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148 information, please see Santos et al., 2015b). A total of 20 sea turtles (mean CCL: 38.1; SD:
149 ±9.5; range: 30–67 cm) distributed throughout all the studied areas ingested cephalopods
150 (Table 1). We found no significant difference in size between sea turtles with and without
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151 squid remains in their intestinal tract (Mann-Whitney test; p = 0.191). Therefore, we do not
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152 expect that size influences plastic ingestion.

153 Nine cephalopod taxa were identified from sixteen pairs and two single beaks found
154 in the diet analysis (Table 1). The most common taxa were Chiroteuthis spp. (7 individuals),
155 followed by Histioteuthis spp. (6 individuals). Our data provide, as far as we know, new
156 records of some cephalopod taxa as prey items of green sea turtles, Chelonia mydas,
157 including the families Argonautidae, Ancistrocheiridae, Cranchiidae, and Octopoteuthidae
158 and the suborder Oegopsida.

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159 Plastic ingestion was recorded for 68.1% (201 individuals) of the 295 green turtles.
160 We retrieved 8251 debris items from intestinal tracts, with a mean of 41 items per turtle that
161 ingested debris (SD: 95.3; range: 1–842 items). A total mass of 633.8 g was recorded, with a
162 mean weight of 3.3 g (SD: 16.5; range: <0.1–216.1 g) per turtle (for detailed information, see
163 Table S2).

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165 Is squid ingestion by sea turtles a result of scavenging behavior?

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166

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167 We argue that the most parsimonious explanation for cephalopod ingestion by
168 juvenile sea turtles is opportunistic scavenging behavior. This conclusion is based on the
169 following: (1) the ingested cephalopods were too large to be captured alive by green turtles

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170 (e.g., Megalocranchia maxima) (Figure 1) and (2) cephalopod species identified from the
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171 GIT of turtles are typically caught as bycatch in deep oceanic fisheries (> 150 m depth)
172 (Perez et al., 2004); thus, carcasses may float and remain available to marine scavengers.
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174 Is scavenging behavior correlated with debris ingestion?

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176 All sea turtles that ingested cephalopods also ingested plastic debris. The amount of
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177 marine debris retrieved from the sea turtles that ingested squid ranged from 1 to 842 items
178 (mean: 214,7; SD: ±202.4), and the mean weight of debris ingested was 25.5 g (standard
179 deviation (SD): ±48.6; range: 0.03-216.1 g). From 20 sea turtles that had ingested
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180 cephalopods, it was possible to determine the cause of death of 15 of them (75%). From that
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181 determination, necropsies revealed that 10 individuals (66.7%) of C. mydas died by debris
182 ingestion, which is remarkably higher than the mean debris ingestion death rates reported for
183 C. mydas in Brazil (10.7%) (for detailed information, see Santos et al., 2015a). Individuals
184 that ingested cephalopods presented significantly higher debris amounts (number and weight)
185 in their intestinal tract than individuals that did not ingest cephalopods (Mann-Whitney test; p
186 < 0.01; Figure 2; Table S2). All turtles that ingested squid exceeded the amount of 0.5 g,
187 which was the minimum amount of debris that was recorded as sufficient to cause the death
188 of a juvenile green turtle through the blockage of the digestive tract (Santos et al., 2015a).

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189 As expected, the model showed that foraging areas have an effect on the amount of
190 ingested plastic by sea turtles; however, the strongest positive effect observed was the
191 occurrence of scavenging behavior (β = 2.35920, p = 0.032) (Figure 3). Detailed results are
192 presented in the Supplemental Material. The foraging areas that had a positive effect on the
193 amount of debris ingestion were areas where turtles extensively foraged in the water column
194 (estuarine areas and cold-water rock reefs; for further information, see Electronic

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195 Supplementary Material and Santos et al., 2015b).

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197 DISCUSSION

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199 Scavenging in nature occurs as a primary feeding strategy (obligate scavengers) or as

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200 a means of gaining additional energy from an unexpected resource (opportunistic feeding
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201 strategy) (Fallows et al., 2013; Kane et al., 2017; Ruxton and Houston, 2004). For obligate
202 and opportunistic scavengers, the encounter rate and handling time are essential parameters to
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have success in a scavenging foraging event (Kane et al., 2017). In the marine realm, carrion
204 can be consumed by any size animal, from small invertebrates to large vertebrates
205 (Colombini and Chelazzi, 2003; Fallows et al., 2013), thus shaping the trophic ecology and
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206 community structure in oceanic and coastal environments (Britton and Morton, 1994; Fallows
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207 et al., 2013; Ruxton and Houston, 2004; Wilson and Wolkovich, 2011).

208 Terrestrial and marine scavenging behaviors are opportunistic strategies performed
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209 mainly by carnivorous animals (Kane et al., 2017). However, herbivores can display
210 noteworthy trophic plasticity in consuming animal matter (Castilla et al., 2011). Green turtles
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211 are an example of an herbivorous species that occasionally forage on animal matter
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212 (Amorocho and Reina, 2007; Lemons et al., 2011; Santos et al., 2015b). The use of animal
213 matter by green turtles is evident during their surface-pelagic phase (i.e., “lost years”), a
214 period when early juveniles remain in surface-pelagic ocean waters before their recruitment
215 to the neritic zone (Witherington et al., 2012), and some studies have pointed out that this diet
216 may persist during the first years after recruitment (Cardona et al., 2009; Reich et al., 2007).
217 Additionally, animal matter seems to be the main diet in some regions of the most southern
218 distribution area of the species (González Carman et al., 2013; Santos et al., 2015b).

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220 Is squid ingestion by sea turtles a result of scavenging behavior?

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222 Scavenging on squid carcasses is widely reported in the marine environment (Cooper
223 et al., 1992; Croxall and Prince, 1996; Lipinski and Jackson, 1989). Natural mortality after
224 spawning events produces larger amounts of floating squid carcasses, and many species of

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225 fish and seabirds benefit from the carcasses (Xavier and Croxall, 2007). The consumption of
226 squid by green turtles has also been reported (Seminoff et al., 2002; Morais et al., 2012). In

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227 this context, Morais et al. (2012) suggested two scenarios to explain the ingestion of squid by
228 green turtles: predatory and scavenging behavior. However, they did not present a clear

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229 conclusion for which was more likely. Our study shed light on this question, and our
230 evidence suggests that the ingestion of cephalopods is better explained by scavenging
231 behavior than by active pursuit predation. Scavenging behavior is suggested based on the
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estimated size of the ingested squids and their association with deep waters (Perez et al.,
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233 2004), both of which are incompatible with the size of juvenile sea turtles and their feeding
234 behavior. Additionally, all the turtles showed evidence of coastal benthic forage, such as the
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235 presence of seagrass and algae in their gastrointestinal tract, which is not compatible with
236 squid habitats (most species were from deep oceanic waters). We are aware that juveniles of
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C. mydas may make movements from nearshore to offshore sites (Williard et al., 2017),
238 although we have no evidence of this from our study or other studies (Santos et al., 2015b),
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239 which may suggest the occurrence of feeding movements from coastal to oceanic waters in
240 our study area.
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242 Is scavenging behavior correlated with debris ingestion?

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244 Opportunistic scavenging behavior provides a suitable energetic trade-off in marine

245 environments (Fallows et al., 2013); however, human-induced environmental changes may
246 turn this adaptive behavior into a maladaptive one, reducing individual fitness (Sih et al.,
247 2011; Tuomainen and Candolin, 2011). In this context, we suggest that the opportunistic
248 scavenging behavior, once a low-cost strategy to acquire protein-rich resources in most ocean
249 waters, became an evolutionary trap (Robertson et al., 2013; Schlaepfer et al., 2002) due to
250 the ubiquity of plastic in the marine realm. Currently, the majority of floating material on the

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251 ocean surface is plastic debris and not animal carcasses (Cózar et al., 2014; Law et al., 2010);
252 hence, the opportunistic strategy to forage on floating material can be lethal to marine
253 animals. Although the foraging area is an important factor influencing elevated plastic
254 ingestion, which is mainly related to variation in debris availability (Barnes et al., 2009) and
255 turtle foraging strategies (Santos et al., 2015a, 2015b), our results showed that scavenging
256 behavior is also an important driver. The mean amount of debris that was ingested by turtles

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257 in this study, possibly through opportunistic scavenging behavior, exceeded by 50-fold the
258 minimum amount of ingested debris that was recorded as sufficient to cause death through

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259 the blockage of the digestive tract (Santos et al., 2015a). Additionally, 66.7% of all turtles
260 that ingested squid died due to debris ingestion through the blockage of the digestive tract, a

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261 number much higher than the highest value found among all studied areas along the Brazilian
262 coast (29%; Santos et al., 2015a). Therefore, the opportunistic scavenging behavior of green
263 turtles led to high rates and loads of plastic ingestion, enhancing their chances of dying and
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being affected by sublethal effects, such as a decrease in nutritional gain and exposure to
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265 chemicals leaching from plastic (Gregory, 2009).

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267 CONCLUSIONS
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269 The relatively recent presence of plastics in society and as a source of pollution in the
270 oceans (Thompson et al., 2009) relative to the evolutionary history of species led us to
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271 contend that common and formerly beneficial scavenging behaviors may now be an
272 important pathway for plastic debris ingestion. The high prevalence of scavenging behavior
273 and its importance to aquatic food webs (Wilson and Wolkovich, 2011) contributed to our
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274 suggestion that scavenging behavior may act as a pathway for plastic debris ingestion,
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275 creating a worrying scenario regarding the rapid spread of plastics throughout all aquatic food
276 webs by either direct or indirect (i.e., ingesting prey that contains plastic) plastic ingestion.
277 Therefore, the study findings may help to explain the causes of widespread plastic ingestion
278 by marine animals with different diets and foraging strategies worldwide. Although, we
279 showed that scavenging behavior can increase plastic ingestion chance in marine
280 environment, it cannot be seeing as the main factor influencing the plastic ingestion to most
281 species. In fact, plastic ingestion seems to be related to plentiful factors, such as visual and
282 olfactory cues, foraging strategy among others (e.g. Santos et al. 2015a, 2016; Savoca et al.

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283 2016, 2017; Schuyler et al. 2014). Therefore, further studies testing plastic ingestion
284 susceptibility must consider more species with different foraging strategies and
285 chemosensory mechanisms in order to explore a broad spectrum of this problem.

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287 ACKNOWLEDGMENTS

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289 We thank the TAMAR/ICMBio team for the collection of stranded dead turtles during their
290 intensive coastal monitoring. We thank M. A. Boldrini for performing most of the green

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291 turtle necropsies and L.M.A. Sousa for the drawings. We were partially supported by
292 Fundação de Amparo à Pesquisa e Inovação do Espírito Santo (FAPES). This study was
293 conducted under the Brazilian System of Authorization and Information on Biodiversity-
294 SISBIO, license number # 21802-3.
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450 TABLE AND FIGURE CAPTIONS
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452 Table 1 – Records of cephalopod ingestion by green turtles along the Brazilian coast. Green
453 turtle size classes were calculated using curved carapace length (cm). Site refers to the
454 location where the individual was found, and ML refers to the mantle length (mm) of
455 cephalopods provided by estimations described in the Material and Methods section. NI

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456 denotes unidentified taxa and indicates when it was not possible to estimate the mantle
457 length. NI* = Oegopsida (Suborder).

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458 Figure 1 – Representation of the size of cephalopod genera in relation to Chelonia mydas.

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459 The size of cephalopods was based on the smallest mantle length recorded for each genus that
460 was ingested by green turtles. The green turtle size represents an individual with 35 cm of
461 curved carapace length, which was the mean size of the turtles that ingested cephalopods
462
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464 Figure 2 – Mean (+SE) weight (A) and number (B) of debris found in Chelonia mydas that
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465 had squid remains (Squid-presence) and those that did not have squid remains (Squid-
466 absence) in their intestinal tract. Both comparisons (weight, N = 294; and number, N = 295)
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467 between the presence and absence of squid presented significant differences in debris
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468 ingestion (p < 0.01) via Mann-Whitney tests.

469
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470 Figure 3 – Effects of predictors (scavenging behavior, foraging areas and carapace curve
471 length) on the amount (number of items) of plastic debris ingested by Chelonia mydas.
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472 Coefficients with 95% confidence intervals that do not overlap with zero are shown in blue
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Individual Site Size class (cm) Family Species ML (mm)
CM 1 09˚39’S, 35˚44’W 30-35 Chiroteuthidae Chiroteuthis sp. 97
Chiroteuthidae Chiroteuthis sp. 121.5
Chiroteuthidae Chiroteuthis sp. 123.9
CM 2 10˚24’S, 36˚26’W 30-35 NI* NI NI
CM 3 10˚53’S, 36˚50’W 35-40 NI NI NI
CM 4 10˚53’S, 36˚50’W 35-40 Ancistrocheiridae Ancistrocheirus lesueuri 94.4
CM 5 11˚33’S, 37˚47’W 35-40 NI NI NI

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CM 6 19˚23’S, 40˚04’W 35-40 Chiroteuthidae Chiroteuthis sp. 131.3
CM 7 19˚49’S, 40˚16’W 30-35 Argonautidae Argonauta sp. NI
CM 8 19˚55’S, 40˚24’W 30-35 Histioteuthidae Histioteuthis corona corona 82.3

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CM 9 20˚19’S, 40˚20’W 30-35 NI NI NI
NI NI NI
CM 10 20˚19’S, 40˚20’W 35-40 Histioteuthidae Histioteuthis corona corona 93.1

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CM 11 20˚19’S, 40˚20’W 30-35 Chiroteuthidae Chiroteuthis sp. 119
CM 12 20˚19’S, 40˚20’W 35-40 Cranchiidae Megalocranchia maxima 528.6
Chiroteuthidae Chiroteuthis sp. NI

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CM 13 20˚19’S, 40˚20’W 30-35 Histioteuthidae Histioteuthis corona corona 103.9
Histioteuthidae Histioteuthis sp. 64.1
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Histioteuthidae Histioteuthis sp. 57.5
CM 14 20˚40’S, 40˚29’W 40-45 NI NI NI
CM 15 21˚02’S, 40˚49’W 30-35 NI NI NI
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CM 16 22˚57’S, 42˚01’W 65-70 NI NI NI

CM 17 20˚52’S, 42˚20’W 30-35 Chiroteuthidae Chiroteuthis sp. 116.6
CM 18 23˚26’S, 45˚04’W 40-45 Octopoteuthidae Octopoteuthis sp. NI
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Chiroteuthidae Chiroteuthis veranyi 111.7

Histioteuthidae Histioteuthis sp. 70.8
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CM 19 27˚07’S, 48˚31’W 35-40 NI NI NI

CM 20 26˚46’S, 48˚38’W 35-40 Cranchiidae Liocranchia reinhardti 113.8
NI* NI NI
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HIGHLIGHTS

• Sea turtles seem to be scavenging on dead squids

• Sea turtles scavenging on dead squids show high plastic ingestion
• Scavenging may be an important pathway for plastic debris ingestion
• Scavenging may help to explain the widespread plastic ingestion by marine animals

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