Mar Biol DOI 10.

1007/s00227-011-1836-y

ORIGINAL PAPER

Diet of juvenile green turtles (Chelonia mydas) associating with artisanal shing traps in a subtropical estuary in Brazil
Shany Mayumi Nagaoka Agnaldo Silva Martins Robson Guimaraes dos Santos Monica Maria Pereira Tognella Eurico Cabral de Oliveira Filho Jeffrey A. Seminoff

Received: 1 June 2011 / Accepted: 30 October 2011 Springer-Verlag 2011

Abstract The diet of 50 juvenile green turtles Chelonia mydas live-captured incidentally by xed shing traps between January and June 2009 in Cananeia Estuarine Lagoon complex, Brazil, was studied through analysis of esophageal lavage samples. Green turtles consumed an omnivorous diet, with 18 food components identied and grouped into 4 categories as follows: terrestrial plants, algae, invertebrates, and seagrass. Black mangrove leaves were of the greatest importance to diet. Turtles incidentally get into xed traps probably because these devices are located on mangrove margins, where they forage. The additional foods suggest that green turtles also feed opportunistically on material adhered to the trap structure and/or on items that cross into its interior. Green turtle diet in estuarine environments appears to be determined by the availability of food components, with some selectivity toward items of apparently greater nutritional value.
Communicated by R. Lewison. S. M. Nagaoka (&) A. S. Martins R. G. dos Santos Departamento de Oceanograa e Ecologia, CCHN, Universidade Federal do Esprito Santo, 29075-910 Vitoria, ES, Brazil e-mail: shanynagaoka@yahoo.com.br M. M. P. Tognella Departamento de Ciencias da Saude Biologica e Agrarias, CEUNES, Universidade Federal do Esprito Santo, 29933-415 Sao Mateus, ES, Brazil E. C. de Oliveira Filho Departamento de Botanica, Universidade Federal de Santa Catarina, 88040-900 Florianopolis, SC, Brazil J. A. Seminoff Southwest Fisheries Science Center, National Marine Fisheries Service (NOAA), 3333 N. Torrey Pines Ct., La Jolla, CA 92037, USA

Introduction The green turtle (Chelonia mydas) is threatened throughout its geographic range due to human overexploitation, incidental mortality related to shing activities, and loss of feeding and nesting habitats (Limpus 1995).Understanding feeding habits of this endangered species is essential to its conservation, as diet directly inuences growth rate, and in turn, reproductive output and demography (Balazs 1982; Bjorndal 1982). Green turtles have a particular life history in terms of its feeding ecology. As hatchlings inhabit oceanic areas and have an omnivorous diet (Bjorndal 1997); after pelagic phase lasting several or more years, they recruit to neritic habitats and shift toward a more herbivorous diet (Bjorndal 1997; Hirth 1997; Reich et al. 2007). It is known that green turtles diet is geographically variable, which makes it critical to conduct foraging studies in many feeding grounds, in order to generate biological information to facilitate site-specic management actions (Seminoff et al. 2002; Andre et al. 2005; Lopez-Mendilaharsu et al. 2005). The diet of green turtles may include algae (Ferreira 1968; Bjorndal et al. 1991) or seagrasses (Mortimer 1981; Moran and Bjorndal 2005) as primary constituents, but some populations may target both food resources (Garnett et al. 1985; Ross 1985, BrandGardner et al. 1999). In addition, green turtles in some areas consume large portions of invertebrates (Seminoff et al. 2002; Bugoni et al. 2003, Amorocho and Reina 2007). The differences in diet composition are probably related to local availability of foods (Garnett et al. 1985), although some level of selectivity has been demonstrated (Bjorndal 1985; BrandGardner et al. 1999; Fuentes et al. 2006; Lopez-Mendilaharsu et al. 2005).

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For green turtle populations found in estuaries, mangrove leaves and propagules have also been described as diet components (Pritchard 1971; Pendoley and Fitzpatrick 1999; Limpus and Limpus 2000). Little is known about feeding habits of green turtles in mangrove areas lacking abundant seagrass pastures and high concentrations of algae, despite the wide distribution of these ecosystems on sheltered coastlines of tropical and subtropical regions, and the signicant presence of green turtles within them. Insights about the biology of green turtles in such areas can help inform regional conservation efforts, which work toward reducing green turtlesheries interactions. A particular feature present in many estuarine areas is artisanal shing with xed traps (FAO 2010). Because there is no direct inuence from the surf of the open sea, estuaries are often highly suitable for construction of these shing devices (Oliveira 2007). Furthermore, many estuaries are under strong tidal inuence, and this is a basic requirement to successfully catch sh with these traps (Mendonca 1998). The xed sh trap found in Brazil, especially in Cananeia, is constructed with organic material such as trees, palms, and bamboo removed from nearby forests (Oliveira 2007) (Fig. 1). The length of trap and the distance between the bamboos sticks vary with the season according to the targeted sh species (Mendonca 1998). The incidental capture of sea turtles by xed traps, especially green turtles, has been cited as very common on the US east coast (Epperly et al. 1995; Manseld 2006; Epperly et al. 2007) and in Brazil (Lima 1999; Nagaoka et al. 2008). However, no study has addressed the foraging tendencies of green turtles captured in these devices nor are

there data on foraging behavior in regions where such sh traps are prevalent. The goal of this study was to investigate the feeding habits of green turtles in one of the largest mangroves estuaries along the Brazilian coast. This is the rst study to characterize the diet and the feeding tendencies of green turtles trapped in xed shing gears in an estuarine subtropical environment in the Atlantic. To evaluate the context of our ndings, we compare our results with those from other green turtle studies in similar habitats. We hope that data presented herein can be of use for developing more appropriate management of green turtles in the southeastern Atlantic Ocean.

Materials and methods Study site Cananeia EstuarineLagoon complex is situated on the southern coast of Sao Paulo, southeastern Brazilian coast (25000 25040 S and 47560 W) (Fig. 2). The region represents one of the most important Brazilian ecosystems preserved (Diegues 1987) and recognized by UNESCO in 1999 as Natural World Heritage Site. It is also considered an important feeding and development area for juvenile green turtles (Bondioli et al. 2008). The mean water depth is 6 m, with a semidiurnal tidal regime, and mean tidal amplitude of approximately 0.8 m (Mesquita and Harari 1988). The climate is warm and humid, with mean annual air and seawater temperatures of 21.4 and 23.8C, respectively (Schaeffer-Novelli et al. 1990), and mean annual precipitation of ca. 2,000 mm (Ramos et al. 1980). The salinity distribution varies depending on tide period and river discharge, ranging from 11 to 34 PSU (Bergamo 2000). The circulation system is driven primarily by tidal ow (Miyao et al. 1986). The area is characterized as a large estuary dominated by mangroves; the three most characteristic species are Laguncularia racemosa (white mangrove), Rhizophora mangle (red mangrove), and Avicennia schaueriana (black mangrove) (Schaeffer-Novelli et al. 1990). The grass Spartina alterniora colonizes the intertidal area, constituting a monospecic narrow fringe in front of mangroves (Schaeffer-Novelli et al. 1990). The algae are generally afxed to mangrove trunks, branches, and pneumatophores anchors (Oliveira 1984). Seagrasses are uncommon and have yet to be cataloged in this region. Green turtle capture From January to June 2009, 11 xed sh traps, locally called cercos-xos, were monitored, once or twice per

Fig. 1 Cerco-xo structure (Adapted from Mendonca 1998)

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Mar Biol Fig. 2 Study central area in Cananeia EstuarineLagoon Complex, southeastern Brazil. The letters represent the main capture points of the green turtles analyzed: A Ilha Comprida, B Ilha do Cardoso, C Ilha da Casca, D Barra do Taquari, E Ilha de Cananeia

week (Fig. 2). Turtles were removed with a hand net when surfacing to breath. We recorded date, trap location, and when possible, approximately how many days each turtle remained inside the trap, based on the interval between monitoring. Dead green turtles that had drowned inciden tally in artisanal shery gillnets set in Cananeia estuary were also collected during the study period. Tagging and measurements Each live-captured green turtle was tagged with Inconel tags (No. 681, National Band & Tag Co., USA) in both fore ippers. We measured curved carapace length (CCL; 0.1 cm) and curved carapace width (CCW; 0.1 cm) with a exible tape, and body weight with a spring balance. Individuals with CCL B 108 cm were considered as probable immature turtles based on the mean size of nesting females at Ascencion Island (Hirth 1997), the primary source rookery for this foraging site (Bondioli 2009). Diet sampling and analysis Food samples were collected via esophageal lavage (Forbes and Limpus 1993), which allowed the rapid retrieval of recently ingested food from the lower esophagus via gentle ushing with clean salt water. After sampling, the turtles were released near to the capture site. Individuals found dead were necropsied, and stomach contents were collected. All food material obtained were preserved in a 4% buffered formalin/seawater solution. Diet items were identied to the lowest possible taxonomic level, based on taxonomic keys and microscopic analysis. Unfortunately, stomach contents from dead individuals were analyzed only qualitatively for major food

groups due to posterior lost of samples; species-specic identication was only possible in the algae group. The relative abundance of food items from esophageal contents was determined using the principles of microstereology and a quantication technique adapted from Forbes (1999). Lavage samples were mixed and then placed in the bottom of a Petri dish to be viewed through a stereo microscope. A grid of 16 graticules (1 cm2 and 0.5 cm) was attached below the Petri dish. Each diet components contribution was determined by counting the number of graticule line endpoints over which the food item was observed relative to the total number of intercepts counted for all components combined. Total volume obtained by water displacement was multiplied by the percentage of relative abundance, resulting in the volume estimated of each item in each sample. Relative volume (%V, Eq. 1) and the frequency of occurrence (%F, Eq. 2) of each food item were calculated as follows. %V Total volume of diet item in all samples ml 100 Total volume of all samples ml 1 %F Number of samples cointaining diet item 100 Total number of samples 2

Statistical analysis MannWhitney U and ANOVA tests were used to evaluate the differences in CCL from individuals captured at different sites. To assess the diet differences according to the capture sites and the in trap time, the individuals were separated into three classes based on trap days: 03 days, 47 days, and [7 days, and a similarity analysis ANOSIM, where R symbolizes dissimilarity between samples, was performed.

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The relative importance of each food item was assessed using the simple resultant index (Rs) (Eq. 3) and the weighted resultant index (Rw) (Eq. 4) (Mohan and Sankaran 1988). These indices were plotted against the corresponding values of the angle h (Figs. 3, 4). A representative food item with similar values of V and F showed h values around 45. The correction factor Q took into account the deviation of h from mid-way, while grading the food items in the order of importance. Both indices have been used in sea turtles diet studies (Lopez-Mendilaharsu et al. 2005, 2008). To evaluate whether the turtles diet was inuenced by the time they were conned in the trap, the Rw for the main food groups consumed in each class was calculated. Rs and Rw were calculated using volume (V) and frequency of occurence (F) (see above) according to the following: 2 1=2 V F2 Rs 3 1=2 100 R V2 F2 1=2 Q V2 F2 Rw 4 1=2 100 RQ V2 + F2 where, Q 45 jh 45j 45 and h tan1 V/F

100

Rwa

Rwb

Rwc
1 1

Resultant Indices

80 60 40 20 0 0
3 3 2 2 1

10

15

20

25

30

35

40

Angle (degrees)
Fig. 4 Resulting index (Rw) in relation to the angle (h) for the main food groups consumed by green turtles in Cananeia estuary, southeastern Brazil, according to time they were conned in xed traps (a = 03 days, b = 47 days and c = [7 days). Food groups in decreasing order of importance: (1) a, b = terrestrial angiosperms (2) a, b = algae, (3) a, b = invertebrates

Results Fifty-three esophageal samples were collected from a total of 50 live-captured green turtles extracted from xed traps.

35

Rw
30

Rs

25 20 15 10
2

5 0 0 5

3 4 5 6

10

15

20

25

Angle (degrees)
Fig. 3 Resulting index (Rw and Rs) in relation to angle (h) for food items found in 53 esophageal contents of 50 green turtles in Cananeia estuary, southeastern Brazil. Food items in decreasing order of importance (with Rw [ 1): (1) Avicennia schaueriana, (2) unidentied terrestrial plant, (3) Spartina alterniora, (4) Ulva spp., (5) Bostrychia spp., (6) Gayralia spp.

Two individuals were recaptured once and twice, respectively. We also analyzed stomach contents from eight dead green turtles that drowned in gillnets. All turtles were considered immature, with mean CCL of 36.9 cm (SD = 3.70, minmax = 31.050.0 cm). There was no effect of capture site on the size of live turtles. There was no difference in diet composition of turtles captured at the various sh-trap sites within the estuary (R = 0.089 and P = 1.8%). The total volume of all esophageal samples combined was 45.9 ml (mean volume per sample = 0.9 1.68 ml). The primary diet species included terrestrial plants (35.1%), algae (20.1%), invertebrates (7.8%), and seagrass (2.4%). Animal items were considered as invertebrates, despite the presence of sh scales and bones that did not present signicant importance to green turtles diet in this study (Table 1). Wood fragments, roots, seeds, substrates, and debris were also found and grouped as other items (34.4%). Additional components were present in trace levels: a red alga from the Delesseriaceae family, colonial diatoms (Chrysophyta), and cyanobacteria Lyngbia sp. With respect to terrestrial plants, leaves of black mangrove Avicennia schaueriana had the largest relative sample volume (19.2%), followed by an unidentied monocot species (Liliopsida class, 8.3%), and nally by the grass Spartina alterniora (7.7%). One individual exclusively consumed amounts of A. schaueriana (sample volume = 0.7 ml). The largest contribution of algae volume was Chlorophyta (13.4%), followed by Rhodophyta (6.4%). Despite the larger volume of green algae, there was a greater diversity of red algae, including a freshwater species (Compsopogon sp.). The red algae Bostrychia radicans, Bostrychia scorpioides, and Bostrychia calliptera were grouped as Bostrychia spp. as it was not possible to distinguish them during the quantication. The green algae Ulva clathrata and another

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Resultant Indices

Mar Biol Table 1 Number of samples containing the food item (n), relative volume (%V), frequency of occurrence (%F), and weighted resultant index (Rw) of the main components of the diet in 53 esophageal contents of 50 green turtles captured by xed shing traps at Cananeia estuary, southeastern Brazil Food item Algae Rhodophyta Bostrychia spp. Caloglossa spp. Compsopogon sp.* Polysiphonia sp. Rhodomelaceae unidentied Chlorophyta Ulva spp. Gayralia spp. Rhizoclonium sp. Chrysophyta Vaucheria sp. Total algae Seagrass Halodule wrightii Terrestrial plants Avicennia schaueriana Spartina alterniora Unidentied monocot Total terrestrial plants Animal items Bryozoa Hydrozoa Eggs of Aplysia sp. Fish (scales and bones) Total animal items Other items * Freshwater algae 4 10 1 11 26 52 0.4 1.0 5.8 0.6 7.8 34.4 7.5 18.9 1.9 20.8 49.1 98.1 0.1 0.3 2.2 24 28 19 39 19.2 7.7 8.3 35.1 45.3 52.8 35.8 73.6 32.8 5.5 6.4 41.6 9 2.4 17.0 0.2 1 30 0.3 20.1 1.9 56.6 14.1 25 7 3 7.6 5.4 0.4 47.2 13.2 5.7 5.3 2.6 14 3 1 2 1 5.6 0.2 0.4 0.1 0.1 26.4 5.7 1.9 3.8 1.9 2.9 n %V %F Rw

Stomach contents analysis Considering the major food groups consumed by the eight dead individuals, terrestrial plants, and algae showed similar frequency of occurrence in the samples (87.5%) followed by invertebrates (50%) and seagrasses (12.5%). Table 2 shows a list of the main algae species found in stomach samples, comparing the %F that these algae species were found in esophageal contents from live-captured individuals. The most representative algae species consumed by live-captured turtles, such as the red and green algae species, were also present in stomach contents of dead turtles. However, the %F of these species consumed by live and dead turtles were different, with only the red alga Bostrychia spp. appearing at approximately the same frequency in both (26.4 and 25%, respectively). The only exception was the presence of a brown alga species (Sargassum sp.) consumed by one dead turtle. Time in xed traps The green turtles studied remained inside the traps from 0 to 10 days, but in the case of nine individuals brought to us by shermen, it was not possible to know how many days they were conned because shermen had not this accurate information. With respect to lavage sample volume, turtles conned for less time (03 days; n = 17) had the greatest mean volume (mean = 1.89 2.67 ml), whereas those that remained for 47 days (n = 19) had the lowest mean volume (mean = 0.25 0.26 ml). Individuals that stayed for more than 7 days (n = 5) presented intermediate volume (mean = 0.81 0.88 ml) in relation to the other two classes of time. There was no difference in diet composition in all classes of connement time (R = -0.017 and P = 62%). Considering the weighted resultant index (Rw), independent of time that turtles remained in traps, the group most abundant and important to diet was terrestrial plants (Fig. 4). Discussion The size of green turtles in the Cananeia LagoonEstuarine complex corresponds to a population composed of juveniles. This is in agreement with the fact that smaller turtles, because they have higher energy demands than adults, prefer sheltered areas where the net energy expenditure during foraging activities is lower than in areas exposed to high energy (Lopez-Mendilaharsu et al. 2005). Using sheltered areas, juvenile turtles are also able to maximize their growth rates via less energy expenditure in combating waves and strong currents (Lopez-Mendilaharsu et al. 2005; Hatase et al. 2006; Amorocho and Reina 2007).

unidentied species of the same genus were also grouped as Ulva spp. The largest contribution to the total volume of invertebrates was gastropod (Aplysia sp.) eggs (5.8%), consumed by only one individual captured in Barra Taquari (Fig. 2, Point D), which remained for up to 4 days in the trap, and comprising 41% of the total esophageal contents collected. According to index weighted (Rw), the food groups consumed according to their ranking in dietary importance were (in decreasing order) as follows: terrestrial plants [ algae [ invertebrates [ seagrass (Table 1). The importance of each food component is shown in Fig. 3, with Avicennia schaueriana (Rw = 32.8) highlighting the greatest importance relative to other diet items.

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Mar Biol Table 2 Comparison of algae species found in stomach contents of eight green turtles drowned in shing nets and the esophageal contents of 50 live-captured green turtles in Cananeia estuary, southeastern Brazil Food item Stomach contents n Rhodophyta Bostrychia spp. Gracilaria sp. Gelidium sp. Murrayella periclados Bryothamnion seaforthii Chondracanthus acicularis Chlorophyta Ulva spp. Gayralia spp. Rhizoclonium sp. Cladophora sp. Phaeophyta Sargassum sp. 1 12.5 1 4 1 1 12.5 50.0 12.5 12.5 17 7 3 32.1 13.2 5.7 Fig. 5 Contribution of major food groups in green turtles diet in different estuarine environments 2 3 2 1 1 1 25.0 37.5 25.0 12.5 12.5 12.5 14 26.4 %F Esophageal contents n %F

n number of individuals and %F frequency of occurrence

Green turtles in Cananeia foraged on a mixed diet dominated by plant material. Despite the limitations of esophageal lavage method in terms of low sample volume, terrestrial plants were clearly the most important diet group followed by algae. According to Bjorndal (1997), green turtles foraging on a mixed diet have lower digestive efciencies, an energetic drawback that likely necessitates greater food consumption rates to offset the low digestibility. In this study, artisanal xed traps along the shores of the study site provided a means to access green turtles for diet study; this method also enabled our evaluation of the rate of interaction between these shing devices and green turtles. Nagaoka et al. (2008) suggested that green turtles entered these sh traps in search of food. However, based on this studythat found low food levels within the sh trapswe believe that green turtles did not voluntarily enter the traps but rather swam into them incidentally, as they are located on mangrove margins, where their feeding habits are found. Although older traps have more encrustations of mollusks, crustaceans, and algae, which may serve as food for green turtles, newly built traps without these organisms also capture turtles, demonstrating that the entrance has no direct relation to searching food associated with traps. Considering that the trap devices are constructed from plant material removed from the local forest (Oliveira 2007) and present a structure that allows the passage of oating material via tidal ushing, the

encrustations occurring on traps, as well as materials that cross into their interior, may be similar to what is present in the surrounding marine environment. The connement time of turtles answered questions related to their behavior during this period in xed traps. When examining the relationship between connement time and volume of samples, it was revealed that volume tends to decrease until a certain residence time in the traps is reached. However, connement for longer periods may result in opportunistic feeding on other materials that are perhaps not a normal diet component for green turtles at this foraging habitat. For example, one turtle conned in a trap at Ilha da Casca (Fig. 2, Point C) for approximately 7 days, was seen eating black mangrove leaves attached to the trap structure, as evidenced by the 100% relative volume of this item in recovered esophageal contents. However, we must clarify that there is no evidences suggesting that all turtles had eaten immediately prior to becoming entrained in the sh traps. We also acknowledge that the low sample volume per turtle may be due to a lack of forage availability within the traps such that all foods had passed the upper esophagus prior to lavage applications. Despite the limitations of sample retrieval due to extended stays within the sh traps, we believe the dietary diversity yielded by these turtles is representative of natural green turtle diet. The stomach contents of eight dead turtles showed that the main food groups are the same as those found in esophageal contents, particularly with respect to algae. The highest frequency of terrestrial plants and algae conrm the importance of both resources to their diet. Pritchard (1971) reported mangrove leaves and roots as part of green turtles diet in the Galapagos Islands, a nding later conrmed by Carrion-Cortez et al. (2010). Pendoley and Fitzpatrick (1999), in reporting their observations of

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green turtles feeding on leaves of Avicennia marina in Western Australia, have highlighted the role of mangrove in green turtle diet. Limpus and Limpus (2000) found the consumption of leaves and cotyledons of A. marina by green turtles feeding in mangroves of Shoalwater Bay, Australia. Diet of green turtles appears to be dictated by the most dominant food items available on the studied area (Fig. 5). Algae were the most important food components in the diet of green turtles along the Pacic coast of the Baja California Peninsula, with food availability dictating its dietary importance in most cased (Lopez-Mendilaharsu et al. 2005, 2008). The same was found by Carrion-Cortez et al. (2010) in Galapagos Islands, Ecuador, where there was abundance of green algae, items that also dominated the diet of green turtles. At Gorgona Island, Colombia, the absence of large algae concentrations and seagrasses and the abundance of animals prey resulted in a largely carnivorous diet by local green turtles (Amorocho and Reina 2007). In contrast, seagrasses were the major diet components of green turtles in Australia (Arthur et al. 2009) and southern Brazil (Guebert 2008) due to their high abundance in these foraging areas. When compared with other green turtle diet studies in Brazil (e.g., Guebert 2008), our results underscore the fact that green turtle diet may vary depending on the local conditions. For example, the area studied by Guebert (2008) on the Parana coast, southern Brazil, is relatively close to the area of the present study, both part of the IguapeCananeiaParanagua EstuarineLagoon complex. The difference in green turtles diet among the site in Guebert (2008) and this study was probably due to the studied area in Parana covering various types of environments (sandy beaches, rocky shores, bays, and mangrove areas), not only exclusively estuarine as the present study, due to diet sampling method adopted (stomach contents analysis). Our study was the rst to describe terrestrial plants as major contributors to green turtle diet. They were represented in decreasing order of importance by the black mangrove Avicennia schaueriana, one unidentied species of monocot plant and Spartina alterniora. The apparent preference of A. schaueriana, although the presence of others mangrove plants in the study area, could be related to palatability of these tissues in detriment of Laguncularia racemosa and Rhizophora mangle. Compared with the others generas, leaves of Avicennia have less cutin, sclereids cells, and tannin contents (Tomlinson 1986). Scholander (1968) describes the genus Avicennia as salt secretors; thus, it probably has lower salt contents in its leaves tissue, which could inuence the palatability. Although it was not possible to identify the leaves of monocot species (class Liliopsida), it is likely coming from

Atlantic forest trees, suggesting that green turtles are opportunistic and might feed on oating materials from rivers carried by the tide. Amorocho and Reina (2007) also described terrestrial plants as sporadic food sources found in convergence currents, suggesting that green turtles in Gorgona complement or replace food items when they are not available. To date, S. alterniora has not been described as an item of relative importance to green turtle diet, with salt marsh environments thus representing an important role in its ecology in this estuary. Although common, algae associated with mangroves trees have not contributed as much in volume in the diet as terrestrial plants, but they do represent the second most consumed food group by green turtles. The presence of wood fragments and substrate in samples is probably due to incidental consumption while turtles were feeding on algae associated with mangroves and substrate. The consumption of Compsopogon sp., a freshwater red alga, although found in only one individual captured in Barra do Taquari (Fig. 2, Point D) may conrm shermen reports in green turtles venturing into rivers for feeding, especially in Taquari river. Algae and seagrass may have provided an alternative food source when available and also a potential nutritional advantage for those turtles able to access areas where they are present (BrandGardner et al. 1999). Despite the prevalence of herbivorous diet items, invertebrates were consumed by 49.1% of green turtles studied (Table 1). This fact is in agreement with Cardona et al. (2009) that states the macrophyte consumption does not begin immediately after recruitment, and animal items remain as part of diet even after starting to feed on herbivorous foods. In southern Brazil, Bugoni et al. (2003) suggests that invertebrates ingestion by green turtles might be incidental in some cases yet selective in others. It is possible that much of the invertebrates consumed by green turtles in Cananeia have been incidental; however, in a specic case of one individual, there was a large amount of sea hare eggs (Aplysia sp.) in its esophageal content, suggesting that it was selective in consuming this item. According to Godley et al. (1998), green turtles select animal items because they provide signicant dietary protein. In conclusion, juvenile green turtles in Cananeia EstuarineLagoon complex have a diet primarily composed of items associated with mangroves, and perhaps they get incidentally entrained in xed traps located along mangrove margins as a result. However, we also have to consider that green turtles might go in close to the mangroves to rest and they get caught in traps during this movement.
Acknowledgments We thank the FAPES and CNPq (ASM grant 308867/2006-8) for the nancial support. This research was permitted by ICMBio/SISBIO (license 18781-1).

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