A Loggerhead Sea Turtle (Caretta caretta) Preying on Fish Within a Mixed-Species Feeding Aggregation

Author(s): Shigetomo Hirama and Blair Witherington Source: Chelonian Conservation and Biology, 11(2):261-265. 2012. Published By: Chelonian Research Foundation DOI: http://dx.doi.org/10.2744/CCB-0918a.1 URL: http://www.bioone.org/doi/full/10.2744/CCB-0918a.1

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Chelonian Conservation and Biology, 2012, 11(2): 261265 g 2012 Chelonian Research Foundation

A Loggerhead Sea Turtle (Caretta caretta) Preying on Fish Within a Mixed-Species Feeding Aggregation SHIGETOMO HIRAMA1 AND BLAIR WITHERINGTON1
1

Florida Fish and Wildlife Conservation Commission, 9700 South A1A, Melbourne Beach, Florida 32951 USA [hirama@cfl.rr.com; witherington@cfl.rr.com]

ABSTRACT. Loggerhead sea turtles (Caretta caretta) feed predominantly on marine crustaceans, mollusks, and other hard-shelled macroinvertebrates. In this note, we report on a subadult loggerhead observed to be feeding on live fish swarming around the turtle in a prey-fish ball. The observation suggests that loggerheads occasionally prey upon certain pelagic fish species other than those encountered as carrion or fishing bait, and may explain accounts of turtles impaled by billfish (Istiophoridae) rostra. Loggerhead sea turtles (Caretta caretta) inhabit tropical, subtropical, and temperate waters worldwide. As they become large immature and adult turtles, loggerheads occupy predominantly benthic habitats in neritic waters (Bolten 2003). In Florida (USA) loggerheads forage within coastal inlets (Butler et al. 1987), lagoons (Ehrhart et al. 2007), bays (Lutcavage and Musick 1985; Schroeder and Foley 1995), and on deepwater (. 10-mdeep) offshore reefs (Girard et al. 2009) in the Gulf of Mexico and Atlantic Ocean. Due to difficulties associated with offshore research, loggerhead foraging in habitats distant from land is not well studied relative to foraging in coastal habitats. When loggerhead diet has been described in the literature, the information has come principally from analyses of gut contents in shore-stranded turtles. Gut contents of stranded loggerheads have included predominantly marine crustaceans, mollusks, and other hard-shelled macroinvertebrates (Lutcavage and Musick 1985; Dodd 1988; Ruckdeschel and Shoop 1988; Burke and Standora 1993; Plotkin et al. 1993; Lazar et al. 2010). In addition to these naturally occurring prey items, longline fishing baits (Revelles et al. 2007) and discarded fish (White 2004) have also been recorded from loggerhead gut contents. Given their apparent preference for slow-moving prey and their limited capacity for rapid movement (S. Hirama and B. Witherington, pers. obs.), loggerheads would not be expected to eat live, fast-moving pelagic fishes. However, many pelagic fishes form compact, ballshaped schools that may change their vulnerability to predation (Pitcher and Parrish 1993). A prey-fish ball

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BIOLOGY, Volume 11, Number 2 2012 to predation by seabirds. Seabirds commonly associate with dolphins and tunas, and in oceanic waters these birds may rely on prey fish available at the surface because of these subsurface predators (Ashmole and Ashmole 1967; Au and Pitman 1986). We hypothesize that the prey-fish school we observed received additional antipredator advantages in being sheltered by the loggerhead. The presence of the turtle likely prevented the approach of predators from the side of the school occupied by the turtle, and when at the center of the ball, the loggerhead likely prevented predators from striking through the school. In our observed case, although the prey fish may have benefitted from their proximity to the turtle, a trade-off was predation by the turtle. Direct observations of loggerheads feeding on food items in the wild are rare. Although there are some firsthand records of interactions between loggerheads and their prey (e.g., Sauls and Thompson 1994), most information on loggerhead foraging comes from ex post facto examination of ingested items. One might infer that most fish found in loggerhead digestive tracts were dead when eaten, given that most fish can easily escape the sluggish movements limiting a loggerheads predatory capacity (Shoop and Ruckdeschel 1982; Lutcavage and Musick 1985). For example, Plotkin et al. (1993) inferred that fish parts found in the digestive tract of stranded loggerheads were from fish eaten as carrion because the guts also contained large numbers of the scavenging gastropod Nassarius acutus. Seney and Musick (2007) found Atlantic menhaden (Brevoortia tyrannus), Atlantic croaker (Micropogonias undulatus), striped bass (Morone saxatilis), bluefish (Pomatomus saltatrix), and seatrout (Cynoscion sp.) in digestive tracts of loggerheads. These authors reasoned that most of these fish had been fed upon as they were either alive or freshly dead, having been restrained or killed by commercial fishing nets. A low frequency of scavenging mud snails in the gut contents supported this observation. In contrast, Revelles et al. (2007) were unable to explain the origins of clupeid fish, which were the highest-frequency item in digestive tracts from 19 dead loggerheads in the western Mediterranean. Tomas et al. (2001) found that teleost fishes were the principal constituent within gut contents of 54 loggerheads captured by trawls in the western Mediterranean. The ingested fish included 13 species, including two clupeids (Sardina pilchardusa and Alosa alosa). The authors concluded that the clupeids had been discarded bycatch, but that the short-snouted seahorse (Hippocampus hippocampus), a slower-moving benthic fish also found in some of the turtle stomachs, may have been consumed alive (Tomas et al. 2001). One question about loggerhead feeding behavior concerns the extent to which they are opportunistic feeders or are selective of targeted food items (Dodd 1988). Although we do not present a sample sufficiently large to answer this question, we do offer an observation of opportunistic feeding on live prey fish in which a

typically contains small fish close to the waters surface (Marchal and Lebourges 1996). The schooling of these prey fish is thought to confuse predators and increase prey survivorship (Partridge 1982; Milinski 1984; Magurran and Pitcher 1987). In the present article, we describe a loggerhead sea turtle that was a focal point of a prey-fish ball, allowing the turtle to feed on the constituent fish. Observation Note. On 30 July 2009, at approximately 1900 hrs, we were returning to port after conducting offshore vessel transects for oceanic juvenile sea turtles in the northern Gulf of Mexico. Approximately 26 km from shore, lat 29u259040N, long 85u059440W, in water 18 m deep, we observed a large immature loggerhead (approximately 70-cm straight carapace length) at the surface (Fig. 1). Above the turtle, seabirds in a mixed-species flock were diving and feeding on fish. The seabird species included black terns (Chlidonias niger, Fig. 1D), Audubons shearwaters (Puffinus lherminieri), royal terns (Thalasseus maximus, Fig. 1C), and sandwich terns (Thalasseus sandvicensis, Fig. 1D). As we approached the feeding site, we observed that the seabirds were feeding on fish swimming out from water and onto the loggerheads partially exposed carapace (Fig. 1A). We recorded a series of 114 images with a Canon single lens reflex camera with a 400-mm focal length, f5.6 telephoto lens. Approximately 20 m from the turtle, we entered the water to observe the loggerhead from beneath the surface. Upon reaching the turtle, we recorded 82 images with a Canon Power Shot A710IS camera within an underwater housing. From our observations and recorded images, we noted that the school of prey fish was in a compact spherical ball similar to that described in the literature (Clua and Grosvalet 2001), except that a turtle was at its center (Fig. 2). We observed 6 species of fish near the loggerhead. The prey fish aggregation consisted of 3 species, round scad (Decapterus punctatus; Carangidae) and sardines (Harengula jaguana and Sardinella aurita; Clupeidae). We observed that predator fish, little tunnies (Euthynnus alletteratus), were congregating and striking from below in order to feed on the prey fish at the perimeter of the ball. We also observed five sharksuckers (Echeneis sp.) circling and occasionally attaching to the turtle, and lesser amberjacks (Seriola fasciata) also maneuvering tightly around the turtle. The loggerhead repeatedly made lateral head movements with its mouth held open. On at least 3 occasions, the turtle closed its mouth on a fish that had swum between its jaws, allowing the turtle to crush the fish and swallow it. These consumed fish could not be specifically identified, but were one of the 3 prey species, Decapterus punctatus, Harengula jaguana, and Sardinella aurita. Discussion. Schools of prey fish decrease their surface-area-to-volume ratio by forming a ball swarm, and further limit their exposure to fish predators by swarming near the waters surface. However, this surface concentration made the prey fish we observed susceptible

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Figure 1. A surfacing subadult Caretta caretta, with Decapterus punctatus, Harengula jaguana, Sardinella aurita, and Seriola fasciata swimming close to the turtles carapace (A, B), and Thalasseus maximus (C), Thalasseus sandvicensis, and Chlidonias niger (D) flying over prey fish.

Figure 2. Underwater images of a subadult Caretta caretta with Decapterus punctatus, Harengula jaguana, Sardinella aurita, Echeneis sp., and Euthynnus alletteratus.

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BURKE, V. AND STANDORA, E. 1993. Factors affecting strandings of cold-stunned juvenile Kemps Ridley and loggerhead sea turtles in Long Island, New York. Copeia 4:11361138. BUTLER, R., NELSON, W., AND HENWOOD, T. 1987. A trawl survey method for estimating loggerhead turtle, Caretta caretta, abundance in five eastern Florida channels and inlets. US National Marine Fisheries Service Fishery Bulletin 85(3): 447453. CLUA, E. AND GROSVALET, F. 2001. Mixed-species feeding aggregation of dolphins, large tunas and seabirds in the Azores. Aquatic Living Resources 14:1118. DODD, C. 1988. Synopsis of the Biological Data on the Loggerhead Sea Turtle Caretta caretta (Linnaeus 1758). US Fish and Wildlife Service Biological Report 88(14), 110 pp. EHRHART, L., REDFOOT, W., AND BAGLEY, D. 2007. Marine turtles of the central region of the Indian River Lagoon System, Florida. Florida Scientist 70(4):415434. FRAZIER, J.G., FIERSTINE, H.L., BEAVERS, S.C., ACHAVAL, F., SUGANUMA, H., PITMAN, R.L., YAMAGUCHI, Y., AND PRIGIONI, C.M. 1994. Impalement of marine turtles (Reptilia, Chelonia: Cheloniidae and Dermochelyidae) by billfishes (Osteichthyes, Perciformes: Istiophoridae and Xiphiidae). Environmental Biology of Fish 39:8596. GIRARD, C., TUCKER, A., AND CALMETTES, B. 2009. Post-nesting migrations of loggerhead sea turtles in the Gulf of Mexico: dispersal in highly dynamic conditions. Marine Biology 156(9):18271839. LAZAR, B., GRACAN, R., KATIC, J., ZAVODNIK, D., JAKLIN, A., AND TVRTKOVIC, N. 2010. Loggerhead sea turtles (Caretta caretta) as bioturbators in neritic habitats: an insight through the analysis of benthic molluscs in the diet. Marine Ecology 32: 6574. LUTCAVAGE, M. AND MUSICK, J. 1985. Aspects of the biology of sea turtles in Virginia. Copeia 2:449456. MAGURRAN, A. AND PITCHER, T. 1987. Provenance, shoal size and the sociobiology of predator-evasion behaviour in minnow shoals. Proceedings of Royal Society B229(1257): 439465. MARCHAL, E. AND LEBOURGES, A. 1996. Acoustic evidence for unusual diel behaviour of a mesopelagic fish (Vinciguerria nimbaria) exploited by tuna. Journal of Marine Science 53: 443447. MILINSKI, M. 1984. A predators costs of overcoming the confusion-effect of swarming prey. Animal Behaviour 32(4): 11571162. PARTRIDGE, B. 1982. Structure and function of fish schools. Scientific American 246(6):9099. PITCHER, T.J. AND PARRISH, J.K. 1993. Functions of shoaling behaviour in teleosts. In: Pitcher, T.J. (Ed.). Behaviour of Teleost Fishes. New York: Chapman and Hall, pp. 363 439. PLOTKIN, P., WICKSTEN, M., AND AMOS, A. 1993. Feeding ecology of the loggerhead sea turtle, Caretta caretta, in the northwestern Gulf of Mexico. Marine Biology 115:115. REVELLES, M., CARDONA, L., AGUILAR, A., AND FERNANDEZ, G. 2007. The diet of pelagic loggerhead sea turtles (Caretta caretta) off the Balearic archipelago (western Mediterranean): relevance of long-line baits. Journal of the Marine Biological Association of the United Kingdom 87:805813. RUCKDESCHEL, C. AND SHOOP, C. 1988. Gut contents of loggerheads: findings, problems, and new questions. In: Schroeder, B.A. (Comp.). Proceedings of the Eighth Annual Workshop on Sea Turtle Biology and Conservation. NOAA Technical Memorandum NMFS-SEFC-214:9798.

loggerhead exhibited predatory behavior that would not seem to be effective in feeding on its more commonly encountered benthic prey species. In the loggerhead we observed, success in capturing prey fish seemed to be linked to the swarming tactic used by the prey fish, a tactic presumed to reduce predation by larger fish. We propose prey-ball feeding as an explanation for the occurrence of tightly schooling pelagic fish in loggerhead diet samples. It remains unclear how commonly prey-fish school around loggerheads, but we point to additional observations that could be explained by these events, such as interactions between billfishes (Istiophoridae) and sea turtles. Frazier et al. (1994) describe four confirmed cases of turtles (loggerhead; olive ridley, Lepidochelys olivacea; leatherback, Dermochelys coriacea; green turtle, Chelonia mydas) being impaled by billfish rostra. In their interpretation, the authors concluded that the interactions may have occurred incidentally due to prey fish (the billfishes target species) congregating around the turtles. We suggest two areas of caution in drawing conclusions from fish found in the gut contents of loggerhead sea turtles. One is that certain pelagic fish species may occasionally be preyed upon while alive and are not always encountered as carrion or fishing bait. A second caution is that researchers are probably correct in identifying other fishes found in loggerhead diet samples to be the result of net-catch, discarded bycatch, or fishing bait, especially if there is additional supporting evidence. ACKNOWLEDGMENTS The observation was made during work funded by grant NA09NMF4720040 from the NOAA, National Marine Fisheries Service (NMFS), and by Floridas Marine Resources Conservation Trust Fund. Work was carried out under a NMFS Permit for Scientific Purposes under the Endangered Species Act (permit 14726). The authors are grateful to D. Adams and E. Matheson for their comments and assistance with fish identification. We thank J. Boyett, B. Crowder, A. Foley, R. Hardy, A. Meylan, and T. OMeara for providing comments on the manuscript. LITERATURE CITED
ASHMOLE, N.P. AND ASHMOLE, M.J. 1967. Comparative feeding ecology of sea birds of a tropical oceanic island. Peabody Museum of Natural History, Yale University, Bulletin 24. New Haven, CT: Peabody Museum of Natural History. AU, D.W.K. AND PITMAN, R.L. 1986. Seabird interactions with dolphins and tuna in the eastern tropical Pacific. Condor 88: 304317. BOLTEN, A. 2003. Variation in sea turtle life history patterns: neritic vs. oceanic developmental stages. In: Lutz, P.L., Musick, J.A., and Wyneken, J. (Eds.). The Biology of Sea Turtles, Volume II. Boca Raton, FL: CRC Press Inc, pp. 243 257.

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SAULS, B. AND THOMPSON, M. 1994. Observations of a loggerhead turtle feeding in the wild. Marine Turtle Newsletter 67:2829. SCHROEDER, B. AND FOLEY, A. 1995. Population studies of marine turtles in Florida Bay. In: Richardson, J.I. and Richardson, T.H. (Comps.). Proceedings of the Twelfth Annual Workshop on Sea Turtle Biology and Conservation; NOAA Technical Memorandum NMFS-SEFSC-361 274, pp. 117 SENEY, E. AND MUSICK, J. 2007. Historical diet analysis of loggerhead sea turtles (Caretta caretta) in Virginia. Copeia 2: 478489. SHOOP, C. AND RUCKDESCHEL, C. 1982. Increasing turtle strandings in the southeast United States: a complicating factor. Biological Conservation 23(3):213215. TOMAS, J., AZNAR, F., AND RAGA, J. 2001. Feeding ecology of the loggerhead turtle Caretta caretta in the western Mediterranean. Journal of Zoology 255:525532. WHITE, M. 2004. Observations of loggerhead turtles feeding on discarded fish catch at Argostoli, Kefalonia. Marine Turtle Newsletter 105:79.
Received: 26 October 2011 Revised and Accepted: 29 December 2011 Handling Editor: Jeffrey A. Seminoff

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