Predicting the Impact of Sea-Level Rise on Caribbean Sea Turtle Nesting Habitat

Abstract:  The projected rise in sea level is likely to increase the vulnerability of coastal zones in the Caribbean, which are already under pressure from a combination of anthropogenic activities and natural processes. One of the major effects will be a loss of beach habitat, which provides nesting sites for endangered sea turtles. To assess the potential impacts of sea-level rise on sea turtle nesting habitat, we used beach profile measurements of turtle nesting beaches on Bonaire, Netherlands Antilles, to develop elevation models of individual beaches in a geographic information system. These models were then used to quantify areas of beach vulnerable to three different scenarios of a rise in sea level. Physical characteristics of the beaches were also recorded and related to beach vulnerability, flooding, and nesting frequency. Beaches varied in physical characteristics and therefore in their vulnerability to flooding. Up to 32% of the total current beach area could be lost with a 0.5-m rise in sea level, with lower, narrower beaches being the most vulnerable. Vulnerability varied with land use adjacent to the beach. These predictions about loss of nesting habitat have important implications for turtle populations in the region.     

The nesting of the Pacific ridley turtle Lepidochelys olivacea on Playa Nancite,Costa Rica

A study of the nesting of the Pacific ridley turtle Lepidochelys olivacea was undertaken in Costa Rica on Playa Nancite — one of two beaches on which the authors had, the previous year, discovered large nesting aggregations of this species. During the 3 1/2-month period of this study, approximately 288,000 turtles nested on the 1300 m-long beach. About 99% of these nested during 3 periods of mass-nesting (arribadas). The predictability of the arribadas was investigated by observing and recording environmental parameters with which they may be correlated. The mass-nesting phenomenon, aspects of nesting behavior, and the adaptive advantages of mass-nesting are discussed. During the period of study, nearly 2,000 turtles were tagged, 102 of which were subsequently recovered. Carapace lengths and widths of 251 tagged turtles were measured. The emergence of hatchlings was recorded daily and, from these data, the incubation period and an indication of egg and hatchling survival was obtained. The activities of the various predators on eggs, hatchlings, and adults are described.     

Breeding biology of the green turtle, Chelonia mydas (Reptilia: Cheloniidae), on Wan-An Island, PengHu archipelago. II. Nest site selection

Nest site selection of the green turtles on Wan-An Island in the summer of 1996 was determined. Turtles (Chelonia mydas) laid on average one clutch for every three emergences. Even though the total track length was 115 m on average, individual lengths varied considerably depending on the nesting beach where the turtles emerged. Limited accessibility, i.e. adequate distance from the nearest village and a well-protected environment, make beaches A and D suitable nesting beaches for green turtles on Wan-An Island. Both total track and nesting track apexes were found clustered in the interface zone, and turtles preferred to reach the vegetation zone once they emerged from the sea. It is suggested that the turtles on Wan-An Island exhibit nest site selection behavior. Based on these results and the high nest site fidelity to their first nesting beach, conservation recommendations are proposed to the county and central governments for the preservation of nesting beaches in their natural state, by prohibiting illegal sand mining and properly controlling turtle watch groups on Wan-An Island. Received: 21 November 1997 / Accepted: 24 December 1998     

Modelling the fate of marine turtle hatchlings

The SLIM oceanographic model was used to examine the fate of hatchling flatback turtles (Natator depressus) in the first two weeks of their dispersal starting at Wild Duck Island, a major turtle-nesting site in the central Great Barrier Reef region of Australia. We ran simulations to investigate the effects of spring versus neap tides, hatchling's swimming behaviour during their first three days at sea, and the location of nesting beaches. The model predicted that up to 50% of the turtle hatchlings entering the sea from the windward, southern beach remained after 14 days in shallow, nearshore waters, irrespective of tides and swimming. These waters are turbid and may be inhospitable to hatchlings. In contrast >80% of hatchlings dispersing from the leeward, northern beach were dispersed to deeper water (10-40 m) in a quasi-stationary dispersion core centred around 10 km north of the nesting beach after 14 days and the offshore spread of the turtle plume was enhanced by the hatchling's seaward swimming during the first three days. This was due to the presence of a coastal boundary layer and a stagnation zone around the northern side of island, but not the southern side. The model confirmed that dispersal from eastern Queensland flatback turtle rookeries is restricted to the lagoons and coastal waters, and that water circulation and hatchling's swimming control dispersion. The model explains why more turtles nest on the northern than the southern side. This study highlights the usefulness of oceanographic models to increase knowledge about a cryptic life stage of marine turtles.     

Chemical aspects of mass spawning in corals. II. (-)-Epi-thunbergol,the sperm attractant in the eggs of the soft coral Lobophytum crassum (Cnidaria: Octocorallia)

Sea turtles migrate between feeding and nesting areas that are often geographically separated by hundreds or thousands of kilometers. Observations of their aggregations at sea and at nesting beaches have led to the hypothesis that sea turtles migrate in socially structured groups. While this migratory strategy is common to many marine vertebrates, socially facilitated behavior is not well documented in testudines. In 1990 and 1991, we attached satellite transmitters to olive ridleys (Lepidochelys olivacea Eschscholtz) found ovipositing together during a mass nesting at Nancite Beach, Costa Rica, to determine whether they migrate independently or in groups after they leave the nesting beach. Results showed that the turtles were not spatially associated during the internesting period, were capable of re-establishing themselves as a group during a subsequent nesting emergence, and were not spatially associated during their postnesting migrations to oceanic feeding areas. We suggested that what appear to be socially structured groups of L. olivacea are in fact individual turtles simultaneously using the same habitat.     

Migratory and reproductive movements of male leatherback turtles (<Emphasis Type="Italic">Dermochelys coriacea</Emphasis>)

The biology of the endangered leatherback turtle (Dermochelys coriacea) at sea is poorly understood. As research has been almost exclusively limited to studies of nesting females, the movements of male turtles and the timing and location of mating activity have remained unknown. We report on the first deployments of satellite tags on male leatherbacks. Male migration to and residency in waters adjacent low-latitude nesting beaches in the western Atlantic suggest that this is where mating occurs, and return migration to these areas reveals male fidelity for breeding sites. Rate of travel decreased markedly after arriving in coastal breeding areas, where males remained for up to 96 days before assuming northward migrations. The initiation of these northward migrations coincided with peak nesting activity in adjacent nesting colonies. Data from satellite-linked time-depth recorders attached to two males revealed diel dive patterns in breeding areas and marked differences in diving behaviour between migratory and breeding periods in one turtle. When male turtles were in waters adjacent nesting colonies, their movements differed from those reported for nesting females, with females ranging farther from shore. Our results suggest that male leatherbacks may be vulnerable to entanglement in coastal fishing gear in waters adjacent nesting beaches.Communicated by R.J. Thompson, St. Johns     

Are anthropogenic factors affecting nesting habitat of sea turtles? The case of Kanzul beach,Riviera Maya-Tulum (Mexico)

Marine coast modification and human pressure affects many species, including sea turtles. In order to study nine anthropogenic impacts that might affect nesting selection of females, incubation and hatching survival of loggerhead (Caretta caretta) and green turtle (Chelonia mydas), building structures were identified along a 5.2 km beach in Kanzul (Mexico). A high number of hotels and houses (88; 818 rooms), with an average density of 16.6 buildings per kilometer were found. These buildings form a barrier which prevents reaching the beach from inland, resulting in habitat fragmentation. Main pressures were detected during nesting selection (14.19% of turtle nesting attempts interrupted), and low impact were found during incubation (0.77%) and hatching (4.7%). There were three impacts defined as high: beach furniture that blocks out the movement of hatchlings or females, direct pressure by tourists, and artificial beachfront lighting that can potentially mislead hatchlings or females. High impacted areas showed lowest values in nesting selection and hatching success. Based on our results, we suggest management strategies to need to be implemented to reduce human pressure and to avoid nesting habitat loss of loggerhead and green turtle in Kanzul, Mexico.     

Some distinguishing characteristics of nesting beaches of the green turtle Chelonia mydas on North West Cape Peninsula,Western Australia

On an 85 km stretch of coastline along the western and northwestern edge of North West Cape Peninsula, Western Australia, are numerous beaches used for nesting by the green turtle Chelonia mydas. Many other beaches in the area are not so used. Nesting beaches displayed three characteristics that distinguished them from non-nesting beaches: the salinity of the sand moisture at nesting depth was lower, the salt content of surface sand was lower, and the beaches were sheltered from prevailing winds. Several beaches on which turtles did not nest exhibited these characteristics, but possessed sand platforms of reduced elevation above sea level. These observations are discussed in relation to the question of what cues green turtles use in selecting nesting beaches.     

Migration of green turtles <Emphasis Type="Italic">Chelonia mydas</Emphasis> from Tortuguero,Costa Rica

During 1955–2003, flipper tags were attached to 46,983 green turtles and ten turtles were fitted with satellite transmitters at Tortuguero, Costa Rica. Eight satellite-tracked turtles stayed within 135 km of the beach and probably returned to nest after release. The internesting area is more extensive than previously documented. Post-nesting migration routes of satellite-tracked turtles varied. Seven turtles swam close to the coast and three turtles swam through oceanic waters before moving toward nearshore areas. Sea surface height anomaly maps indicate that oceanic movements were consistent with the southwestern Caribbean gyre. Circling and semi-circling turtles could have been disoriented but submergence and surface times suggest they may have been feeding in Sargassum sp. concentrations. Rapid post-nesting migrations (mean 2.2 km hr⁻¹) ended on benthic feeding grounds in shallow waters (<20 m) off Belize (n=1), Honduras (n=1) and Nicaragua (n=8). The spatial distribution of migration end points (n=10) and tag returns (n=4,669) are similar. Fishermen in Nicaragua target green turtles along migratory corridors and on foraging grounds. Management efforts are urgently needed in Nicaragua, particularly in the high-density feeding areas south and east of the Witties (N14°09 W82°45). The proximity of foraging grounds to the nesting beach (mean 512 km) may permit female turtles to invest more energy in reproduction and hence the Tortuguero population may have greater potential for recovery than other green turtle nesting populations. Recovery of the Tortuguero green turtle population will benefit countries and marine ecosystems throughout the Caribbean, especially Nicaragua.     

Dune vegetation fertilization by nesting sea turtles

Sea turtle nesting presents a potential pathway to subsidize nutrient-poor dune ecosystems, which provide the nesting habitat for sea turtles. To assess whether this positive feedback between dune plants and turtle nests exists, we measured N concentration and delta15N values in dune soils, leaves from a common dune plant (sea oats [Uniola paniculata]), and addled eggs of loggerhead (Caretta caretta) and green turtles (Chelonia mydas) across a nesting gradient (200-1050 nests/km) along a 40.5-km stretch of beach in east central Florida, USA. The delta15N levels were higher in loggerhead than green turtle eggs, denoting the higher trophic level of loggerhead turtles. Soil N concentration and delta15N values were both positively correlated to turtle nest density. Sea oat leaf tissue delta15N was also positively correlated to nest density, indicating an increased use of augmented marine-based nutrient sources. Foliar N concentration was correlated with delta15N, suggesting that increased nutrient availability from this biogenic vector may enhance the vigor of dune vegetation, promoting dune stabilization and preserving sea turtle nesting habitat.     

Satellite tracking of migrating loggerhead sea turtles (<Emphasis Type="Italic">Caretta caretta</Emphasis>) displaced in the open sea

Loggerhead turtles (Caretta caretta) are known to migrate towards fixed, individually-specific residential feeding grounds. To study their spatial behaviour and their navigational ability, five loggerheads nesting in South Africa were captured when about to start their postnesting migration and tracked by satellite after having been displaced from their usual migratory route. The first turtle, released south of Madagascar about 1,148 km from the capture site, moved west up to mainland Africa and then reached her feeding grounds by following the coast. A second turtle, released farther away (2,140 km) close to La Réunion Island, stopped for some time on the Madagascar east coast, then turned southwards to round the island and regain the African mainland in the northwest, without however allowing us to establish the location of her residential grounds. Three other turtles were released off the Tanzanian coast, 2,193 km north of their nesting area, at the northern edge of the distribution of the feeding grounds along the African coast. All of them headed north, and one turtle found her residential grounds located north of the release site. The other two females started long-distance oceanic wanderings in which they crossed nearly the entire Indian Ocean, apparently being transported by the sea currents of the region. We conclude that adult loggerhead turtles are apparently unable to compensate for the displacement and can return to a pelagic life style characteristic of juvenile turtles. These findings suggest that South African loggerheads rely on simple orientation mechanisms, such as the use of the coastline, as a guide, and compass orientation, possibly integrated by spatiotemporal programmes and/or acquired maps of familiar sites.Communicated by R. Cattaneo-Vietti, Genova     

Population structure and genetic diversity in green turtles nesting at Tortuguero,Costa Rica,based on mitochondrial DNA control region sequences

The green turtle (Chelonia mydas) nesting population at Tortuguero, Costa Rica, is the largest nesting aggregation in the Atlantic, by at least an order of magnitude. Previous mitochondrial DNA (mtDNA) surveys based on limited sampling (n = 41) indicated low genetic diversity and low gene flow with other Caribbean nesting colonies. Furthermore, a survey of nuclear DNA diversity invoked the possibility of substructure within the Tortuguero rookery. To evaluate these characteristics, mtDNA control region sequences were determined for green turtles nesting at Tortuguero in 2001 (n = 157) and 2002 (n = 235). The increased sample revealed three additional haplotypes; five haplotypes are now known for Tortuguero female green turtles. Analyses of molecular variance indicated that there was no significant spatial population structure along the 30-km nesting beach. In addition, no temporal population structure was detected either between the two nesting seasons or within the nesting season. As a result of the larger sample size and additional haplotypes, estimates of genetic separation among Caribbean nesting colonies have changed and the concordance of phylogenetic and phylogeographic patterns reported in the past for green turtles in the Greater Caribbean has weakened. The five haplotypes from Tortuguero represent 36% of the haplotypes identified in green turtle nesting aggregations in the Greater Caribbean and 17% of the haplotypes known to occur in nesting or foraging aggregations in the Greater Caribbean. Haplotype diversity (0.16) and nucleotide diversity (0.0034) for the Tortuguero population are substantially lower than those for the combined rookeries in the Greater Caribbean (0.44 and 0.0078, respectively). Although comprehensive evaluation of regional genetic diversity requires nuclear DNA data, our study indicates that conserving genetic diversity in Caribbean green turtles will require careful management of the smaller rookeries in addition to the Tortuguero rookery.     

Migratory activity by hatchling loggerhead sea turtles (Caretta caretta L.): evidence for divergence between nesting groups

The South Florida subpopulation of loggerhead sea turtles (Caretta caretta L.) nests with great fidelity on either the southeast or the southwest coast of Florida, USA. The hatchlings that emerge from those nests must swim in opposite directions and search for different surface currents to migrate away from continental shelf waters. In this laboratory study, we compared the pattern of swimming activity shown by the hatchlings from each coast over the first 6 days of migration. Turtles from both coasts were equally active during their “frenzy” period (the first 24 h of swimming) and during the daylight hours of the 5 days that followed (the “postfrenzy” period). However, the west coast turtles were significantly more active than the east coast turtles during the nocturnal portion of the postfrenzy period. This difference may be related to the greater distance southwest coast turtles must negotiate to locate surface currents for transport out of the Gulf of Mexico and into the Atlantic Ocean basin. These differing behavioral strategies may be genetically determined, as similar correspondence between activity and distance is well known among migratory populations of birds and fish and is often based upon inherited programs of endogenously driven activity. Alternatively, behavioral differences between the two nesting groups could be a manifestation of phenotypic plasticity that arises as the hatchlings respond to unique environmental cues on each coast.     

Migratory activity by hatchling loggerhead sea turtles (Caretta caretta L.): evidence for divergence between nesting groups

The South Florida subpopulation of loggerhead sea turtles (Caretta caretta L.) nests with great fidelity on either the southeast or the southwest coast of Florida, USA. The hatchlings that emerge from those nests must swim in opposite directions and search for different surface currents to migrate away from continental shelf waters. In this laboratory study, we compared the pattern of swimming activity shown by the hatchlings from each coast over the first 6 days of migration. Turtles from both coasts were equally active during their “frenzy” period (the first 24 h of swimming) and during the daylight hours of the 5 days that followed (the “postfrenzy” period). However, the west coast turtles were significantly more active than the east coast turtles during the nocturnal portion of the postfrenzy period. This difference may be related to the greater distance southwest coast turtles must negotiate to locate surface currents for transport out of the Gulf of Mexico and into the Atlantic Ocean basin. These differing behavioral strategies may be genetically determined, as similar correspondence between activity and distance is well known among migratory populations of birds and fish and is often based upon inherited programs of endogenously driven activity. Alternatively, behavioral differences between the two nesting groups could be a manifestation of phenotypic plasticity that arises as the hatchlings respond to unique environmental cues on each coast.     

Magnetic orientation by hatchling loggerhead sea turtles (Caretta caretta) from the Gulf of Mexico

Loggerhead sea turtle hatchlings emerge from nests on either the east or west coast of the South Florida peninsula and then migrate offshore in opposite directions. Under laboratory conditions, magnetic cues induce east coast hatchlings to swim in directions that promote their transport by oceanic surface currents, such as the North Atlantic gyre. However, the surface currents used by west coast hatchlings are unknown. We examined the responses of west (Sarasota) hatchlings to magnetic cues in the Gulf of Mexico, the Florida Straits, and the Gulf Stream to determine their (1) likely migratory routes (2) orientation where currents lead into the Atlantic Ocean, and (3) orientation adjacent to Florida’s east coast. The results suggest that migration inside Gulf waters may be circuitous, that the turtles respond appropriately to enter Atlantic waters, and that orientation along Florida’s east coast probably promotes transport by the Gulf Stream into the North Atlantic gyre.     

Cost‐Effectiveness of Alternative Conservation Strategies with Application to the Pacific Leatherback Turtle

Although holistic conservation addressing all sources of mortality for endangered species or stocks is the preferred conservation strategy, limited budgets require a criterion to prioritize conservation investments. We compared the cost‐effectiveness of nesting site and at‐sea conservation strategies for Pacific leatherback turtles (Dermochelys coriacea). We sought to determine which conservation strategy or mix of strategies would produce the largest increase in population growth rate per dollar. Alternative strategies included protection of nesters and their eggs at nesting beaches in Indonesia, gear changes, effort restrictions, and caps on turtle takes in the Hawaiian (U.S.A.) longline swordfish fishery, and temporal and area closures in the California (U.S.A.) drift gill net fishery. We used a population model with a biological metric to measure the effects of conservation alternatives. We normalized all effects by cost to prioritize those strategies with the greatest biological effect relative to its economic cost. We used Monte Carlo simulation to address uncertainty in the main variables and to calculate probability distributions for cost‐effectiveness measures. Nesting beach protection was the most cost‐effective means of achieving increases in leatherback populations. This result creates the possibility of noncompensatory bycatch mitigation, where high‐bycatch fisheries invest in protecting nesting beaches. An example of this practice is U.S. processors of longline tuna and California drift gill net fishers that tax themselves to finance low‐cost nesting site protection. Under certain conditions, fisheries interventions, such as technologies that reduce leatherback bycatch without substantially decreasing target species catch, can be cost‐effective. Reducing bycatch in coastal areas where bycatch is high, particularly adjacent to nesting beaches, may be cost‐effective, particularly, if fisheries in the area are small and of little commercial value. Rentabilidad de Estrategias de Conservación Alternativas Aplicadas a Tortugas Laúd del Pacífico     

Origins of green turtle (<Emphasis Type="Italic">Chelonia mydas</Emphasis>) feeding aggregations around Barbados,West Indies

Although green turtles (Chelonia mydas Linnaeus) do not nest in Barbados, the easternmost island in the Caribbean archipelago, juveniles are regularly seen foraging in nearshore waters. To examine the stock composition of this foraging population, mitochondrial (mt) DNA control region sequences were analysed from 60 juvenile (31–70 cm curved carapace length) green turtles and compared with data published for key nesting populations in the Atlantic, as well as other feeding grounds (FGs) in the Caribbean. Eight distinct haplotypes were recognised among the 60 individual green turtles sampled around Barbados. Three of the haplotypes found have only previously been reported from western Caribbean nesting beaches, and two only from South Atlantic beaches. The nesting beach origin of one of the Barbados FG haplotypes is as yet unidentified. Stock mixture analysis based on Bayesian methods showed that the Barbados FG population is a genetically mixed stock consisting of approximately equal contributions from nesting beaches in Ascension Island (25.0%), Aves Island/Surinam (23.0%), Costa Rica (19.0%), and Florida (18.5%), with a lesser but significant contribution from Mexico (10.3%). Linear regression analysis indicated no significant effects of rookery population size or distance of the rookery from the FG on estimated contributions from the source rookeries to the Barbados FG. Our data suggest that the similar-sized green turtles sampled on the Barbados FG are a mixed stock of more diverse origins than any previously sampled feeding aggregations in the Caribbean region. The relatively large contribution from the Ascension Island rookery to the Barbados FG indicates that hatchlings from distant rookeries outside the Caribbean basin enter the North Atlantic gyre and become a significant part of the pool from which eastern Caribbean foraging populations are derived. These data support a life cycle model that incorporates a tendency of immatures to migrate from their initial foraging grounds at settlement towards suitable foraging grounds closer to their natal rookeries as they mature.Communicated by P.W. Sammarco, Chauvin     

Coastal types of graben: the Gulf of Gökova,Mugla-SW Turkey

Grabens are formed under the influence of extensional forces in a normal fault system. The Gulf of Gökova is an active graben located in SW Turkey. Active grabens such as the Gökova enclose highly faulted rocky coasts. Despite the existence of these rocky coasts, examination of google earth images, field studies, DEM analysis and previous studies in the Gulf of Gökova graben revealed that there could be different types of coasts in the region. In this study the factors causing the occurrence and potential environmental effects of the diverse coast types are evaluated. By using the Fairbridge (2004) coastal classification system, the coast types identified in the Gulf of Gökova graben can be classified as; A. Soft-less consolidated-erodible; A1: relatively Insoluble: detrital and loose beach, A2: soluble: beachrock and eolinite B. hard-cliffed-rocky, B1: longevity of hard-rock coast and B2: fault controlled cliffs. The percentages of these classes generally decreased from B1 (79%), A1 (12.4%), B2 (8.3%) and A2 (0.3%) in the study area. As a result of longshore currents, A1-type coasts usually develop as large plain adjacent streams and also in pocket beaches as narrow-long strips near rocky coasts. A2 type is observed in one location within the Gökova region, possibly due to local environmental conditions. However, B1 type developed in peridotites, cherty limestone and cliffs probably because of the active fault system and where the fault plane cut the coast, B2 type occurred. Possible threats to the Gökova region can be attributed to sea level rise owing to tectonism and global warming. It is anticipated that inundation, coastal erosion and salt water intrusion may also affect it.     

Effects of Organized Turtle Watches on Loggerhead (Caretta caretta) Nesting Behavior and Hatchling Production in Florida

To evaluate the effects of organized turtle watches on female sea turtles and their eggs, we quantified nesting behavior and hatchling production of loggerhead turtles ( Caretta caretta ) in south Brevard Country, Florida, U.S.A. We compared the duration of five phases of nesting behavior, the directness of the turtle's return path, rate of travel during return crawl, hatching success, and hatchling emergence success between experimental and control turtles. Experimental turtles nested while observed by an organized turtle watch group consisting of at least 15 people; control turtles were not observed by a turtle watch group. Experimental turtles spent significantly less time camouflaging nest sites than did control turtles. The duration of the other four phases of nesting behavior were not significantly different between the two groups. Experimental turtles also traveled less-direct paths during return crawls, although their rates of travel were not significantly different from those of control turtles. Hatching success and hatchling emergence success were not significantly different between experimental and control turtle nests in either year. Although turtle watch groups influenced nesting behavior, they were not found to be detrimental to hatchling production. Florida's turtle watch program is a means for garnering public support for sea turtle conservation through education, and it should continue.     

Discrimination of ocean wave features by hatchling loggerhead sea turtles, Caretta caretta

At the beginning of their offshore migration, hatchling sea turtles orient directly into oceanic waves as they swim away from land. Recent experiments have demonstrated that hatchlings swimming underwater can determine the propagation direction of waves by monitoring the circular movements they experience as waves pass above. During July and August 1993, we studied how loggerhead sea turtle hatchlings (Caretta caretta L.) from the east coast of Florida, USA, responded to a range of wave parameters. We constructed a wave simulator to reproduce in air the circular movements that normally occur beneath small ocean waves. Hatchlings suspended in air and subjected to these orbital movements attempted to orient into simulated waves when periods and amplitudes were similar to those found near the Florida coast. Orbital movements with longer periods (greater than 10 s), however, failed to elicit responses. The results demonstrate that hatchling loggerheads can distinguish between waves with different periods and amplitudes, and that Florida hatchlings respond most strongly to orbital movements closely resembling those of waves that occur near their natal beach. Received: 28 May 1996 / Accepted: 17 September 1996