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In addition to providing animals with a source of directional or ‘compass’ information, Earth’s magnetic field also provides a potential source of positional or ‘map’ information that animals might exploit to assess location. In less than a generation, the idea that animals use Earth’s magnetic field as a kind of map has gone from a contentious hypothesis to a well-established tenet of animal navigation. Diverse animals ranging from lobsters to birds are now known to use magnetic positional information for a variety of purposes, including staying on track along migratory pathways, adjusting food intake at appropriate points in a migration, remaining within a suitable oceanic region, and navigating toward specific goals. Recent findings also indicate that sea turtles, salmon, and at least some birds imprint on the magnetic field of their natal area when young and use this information to facilitate return as adults, a process that may underlie long-distance natal homing (a.k.a. natal philopatry) in many species. Despite recent progress, much remains to be learned about the organization of magnetic maps, how they develop, and how animals use them in navigation.

During the mass nesting of olive ridley sea turtles, Lepidochelys olivacea, thousands of adult females aggregate offshore, then emerge from the ocean together to lay their eggs along small, specific stretches of coastline. Little is known about the mechanisms that underlie synchronization of mass nesting. As a first step towards identifying environmental variables that control or affect the timing of mass-nesting events, we analysed 10 years of nesting data from Ostional, Costa Rica, a Pacific beach where mass-nesting events occur throughout the year. The onset of mass nesting was not uniformly distributed across lunar phase, with mass nesting occurring predominantly near last-quarter moon. Statistical models were developed to investigate whether environmental variables could be used to predict the timing and size of mass-nesting events. Predictions of the onset of nesting were improved by considering lunar phase and the time since the previous mass-nesting event as well as several oceanographic variables. Predictions of the number of turtles participating in nesting events were improved by considering salinity, relative humidity and nearshore current velocity, three parameters related to weather patterns and rainfall. Overall, the results imply that mass-nesting events at Ostional tend to occur near last-quarter moon but do not follow an invariant lunar or seasonal pattern. Instead, the results suggest that physiological and environmental factors interact to influence mass-nesting events, with no single environmental variable serving as an infallible predictor of either timing or size.

Sea turtles are often restrained manually for brief periods during veterinary evaluation and care in rescue, rehabilitation, research, and aquarium settings. Blood gas values and lactate are routinely evaluated during triage of sea turtles, and lactate clearance is of prognostic significance in cold-stunned individuals. Although increases in blood lactate have been associated with muscle exertion, experimental forced submergence, trawl and pound net capture, and general anesthesia, changes in blood lactate associated with short periods of manual restraint have not been evaluated. Venous blood gas and lactate values were tested in 16 juvenile loggerhead sea turtles (

As the largest and most diverse vertebrate group on the planet, fishes have evolved an impressive array of sensory abilities to overcome the challenges associated with navigating the aquatic realm. Among these, the ability to detect Earth's magnetic field, or magnetoreception, is phylogenetically widespread and used by fish to guide movements over a wide range of spatial scales ranging from local movements to transoceanic migrations. A proliferation of recent studies, particularly in salmonids, has revealed that fish can exploit Earth's magnetic field not only as a source of directional information for maintaining consistent headings, but also as a kind of map for determining location at sea and for returning to natal areas. Despite significant advances, much about magnetoreception in fishes remains enigmatic. How fish detect magnetic fields remains unknown and our understanding of the evolutionary origins of vertebrate magnetoreception would benefit greatly from studies that include a wider array of fish taxa. The rich diversity of life-history characteristics that fishes exhibit, the wide variety of environments they inhabit, and their suitability for manipulative studies, make fishes promising subjects for magnetoreception studies.

Radio frequency electromagnetic noise (RF) of anthropogenic origin has been shown to disrupt magnetic orientation behavior in some animals. Two sources of natural RF might also have the potential to disturb magnetic orientation behavior under some conditions: solar RF and atmospheric RF. In this review, we outline the frequency ranges and electric/magnetic field magnitudes of RF that have been shown to disturb magnetoreceptive behavior in laboratory studies and compare these to the ranges of solar and atmospheric RF. Frequencies shown to be disruptive in laboratory studies range from 0.1 to 10 MHz, with magnetic magnitudes as low as 1 nT reported to have effects. Based on these values, it appears unlikely that solar RF alone routinely disrupts magnetic orientation. In contrast, atmospheric RF does sometimes exceed the levels known to disrupt magnetic orientation in laboratory studies. We provide a reference for when and where atmospheric RF can be expected to reach these levels, as well as a guide for quantifying RF measurements.

Magnetotactic bacteria (MTB) are a diverse group of highly motile Gram-negative microorganisms with the common ability to orient along magnetic field lines, a behavior known as magnetotaxis. Ubiquitous in aquatic sediment environments, MTB are often microaerophilic and abundant at the oxic/anoxic interface. Magnetic field sensing is accomplished using intracellular, membrane-encased, iron-containing minerals known as magnetosomes. The chemistry, morphology and arrangement of magnetosomes differs substantially among different MTB. Although magnetic field sensing mechanisms, genetic bases and protein functions have been elucidated in select model organisms such as the Magnetospirillum strains and Desulfovibrio RS-1, not all findings are applicable to diverse clades of MTB. As the number of identified species has increased, it has become evident that many of the characteristics and mechanisms once presumed to be prototypical of MTB are in fact not universal. Here we present a general overview of the current state of MTB research for readers outside of the realm of prokaryotic research, focusing on recent discoveries, knowledge gaps and future directions. In addition, we report new insights acquired using holographic technology to observe and quantify microbial responses in magnetic fields that are earth-strength or weaker, providing a new ecophysiological approach to in situ MTB research.

Diverse taxa use Earth’s magnetic field in combination with other sensory modalities to accomplish navigation tasks ranging from local homing to long-distance migration across continents and ocean basins. Several animals have the ability to use the inclination or tilt of magnetic field lines as a component of a magnetic compass sense that can be used to maintain migratory headings. In addition, a few animals are able to distinguish among different inclination angles and, in effect, exploit inclination as a surrogate for latitude. Little is known, however, about the role that magnetic inclination plays in guiding long-distance migrations. In this paper, we use an agent-based modelling approach to investigate whether an artificial agent can successfully execute a series of transequatorial migrations by using sequential measurements of magnetic inclination. The agent was tested with multiple navigation strategies in both present-day and reversed magnetic fields. The findings (i) demonstrate that sequential inclination measurements can enable migrations between the northern and southern hemispheres, and (ii) demonstrate that an inclination-based strategy can tolerate a reversed magnetic field, which could be useful in the development of autonomous engineered systems that must be robust to magnetic field changes. The findings also appear to be consistent with the results of some animal navigation experiments, although whether any animal exploits a strategy of using sequential measurements of inclination remains unknown.

Pfaller et al. report that sea turtles respond to odors from biofouled plastic debris with the same behavior that is elicited by food odors, providing a possible unifying explanation for why sea turtles interact with marine plastic.

A variety of animals sense Earth's magnetic field and use it to guide movements over a wide range of spatial scales. Little is known, however, about the mechanisms that underlie magnetic field detection. Among teleost fish, growing evidence suggests that crystals of the mineral magnetite provide the physical basis of the magnetic sense. In this study, juvenile Chinook salmon (Oncorhynchus tshawytscha) were exposed to a brief but strong magnetic pulse capable of altering the magnetic dipole moment of biogenic magnetite. Orientation behaviour of pulsed fish and untreated control fish was then compared in a magnetic coil system under two conditions: (1) the local magnetic field; and (2) a magnetic field that exists near the southern boundary of the natural oceanic range of Chinook salmon. In the local field, no significant difference existed between the orientation of the control and pulsed groups. By contrast, orientation of the two groups was significantly different in the magnetic field from the distant site. These results demonstrate that a magnetic pulse can alter the magnetic orientation behaviour of a fish and are consistent with the hypothesis that salmon have magnetite-based magnetoreception.

Certain long-distance migratory animals, such as salmon and sea turtles, are thought to imprint on the magnetic field of their natal area and to use this information to help them return as adults. Despite a growing body of indirect support for such imprinting, direct experimental evidence thereof remains elusive. Here, using the fruit fly as a magnetoreceptive model organism, we demonstrate that exposure to a specific geographic magnetic field during a critical period of early development affected responses to a matching magnetic field gradient later in life. Specifically, hungry flies that had imprinted on a specific magnetic field from 1 of 3 widely separated geographic locations responded to the imprinted field, but not other magnetic fields, by moving downward, a geotactic behavior associated with foraging. This same behavior occurred spontaneously in the progeny of the next generation: female progeny moved downward in response to the field on which their parents had imprinted, whereas male progeny did so only in the presence of these females. These results represent experimental evidence that organisms can learn and remember a magnetic field to which they were exposed during a critical period of development. Although the function of the behavior is not known, one possibility is that imprinting on the magnetic field of a natal area assists flies and their offspring in recognizing locations likely to be favorable for foraging and reproduction.

The Galapagos sea lion ( Zalophus wollebaeki) is an otariid species endemic to the Galapagos archipelago and is currently listed as endangered. The ocular trematode Philophthalmus zalophi was recently reported to affect the survival of juvenile Galapagos sea lions on Santa Cruz Island, resulting in marked ophthalmic changes. This study evaluated the ophthalmic disease and histopathologic effects of P. zalophi on juvenile Galapagos sea lions in the largest rookery located on San Cristóbal Island. Twenty juvenile Galapagos sea lions (10 male and 10 female) were evaluated among five sites in the rookery El Malecón. Ophthalmic examination, including fluorescein staining and evaluation of the adnexa, cornea, and sclera, were performed on each eye. The presence, number, and location of ocular parasites were determined, and parasites were collected for identification. Conjunctival biopsy was performed on 11 animals: 2 that lacked parasites and gross lesions and 9 with both parasites and gross lesions. All parasites collected were confirmed as P. zalophi and identified in 80% (16/20) of the study animals and 70% (28/40) of the examined eyes. Philophthalmus zalophi was most frequently found attached to the nictitating membrane but also located on the palpebral conjunctiva or cornea. The most common clinical signs were varying degrees of conjunctival hyperemia (28/40 eyes), most frequently of the nictitating membrane and mucoid ocular discharge (12/40 eyes). The number of parasites was significantly associated with the degree of conjunctival hyperemia ( P0.001). Histopathology of conjunctival biopsies revealed organized lymphoid follicles and lymphoplasmacytic infiltrates. The histopathologic changes and gross lesions were likely due to the parasite's attachment to the conjunctiva. This study provides additional details of P. zalophi infection in juvenile Galapagos sea lions. Further research is warranted to detail the life cycle of this parasite, transmission to sea lions, and potential treatment protocols.

Diverse animals use Earth’s magnetic field to guide their movements, but the neural and molecular mechanisms underlying the magnetic sense remain enigmatic. One hypothesis is that particles of the mineral magnetite (Fe3O4) provide the basis of magnetoreception. Here we examined gene expression in the central nervous system of a magnetically sensitive invertebrate, the Caribbean spiny lobster (Panulirus argus), after applying a magnetic pulse known to alter magnetic orientation behavior. Numerous genes were differentially expressed in response to the pulse, including 647 in the brain, 1256 in the subesophageal ganglion, and 712 in the thoracic ganglia. Many such genes encode proteins linked to iron regulation, oxidative stress, and immune response, consistent with possible impacts of a magnetic pulse on magnetite-based magnetoreceptors. Additionally, however, altered expression also occurred for numerous genes with no apparent link to magnetoreception, including genes encoding proteins linked to photoreception, carbohydrate and hormone metabolism, and other physiological processes. Overall, the results are consistent with the magnetite hypothesis of magnetoreception, yet also reveal that in spiny lobsters, a strong pulse altered expression of > 10% of all expressed genes, including many seemingly unrelated to sensory processes. Thus, caution is required when interpreting the effects of magnetic pulses on animal behavior.

OBJECTIVE: To determine the combined mydriatic effects of topical rocuronium bromide and phenylephrine in juvenile loggerhead turtles and identify any adverse effects associated with treatment. ANIMALS STUDIED: Eleven juvenile loggerhead turtles (Caretta caretta). PROCEDURES: Four 20 μL drops of rocuronium bromide and four 20 μL drops of 10% phenylephrine were placed into the right eye at 2‐minute intervals of 5 turtles, while the same volume of saline was administered to six control turtles. A pupilometer recorded pupil measurements at rest and following a light stimulus at 2, 15, 30, 60, 120, 150, 180, 210, 240, 300, and 360 minutes following delivery of the final drop to the ocular surface. Intraocular pressure (IOP) was also measured at similar time points. RESULTS: The nonilluminated and light‐stimulated pupillary diameter of the right eye of treated turtles was significantly greater than baseline starting at 120 and 15 minutes, respectively. Light‐stimulated pupillary diameter of treated eyes was greater than that of control eyes from time 15 minutes until the end of the treatment period. No systemic side effects were noted over a 24 hours period following treatment and all turtles showed normal behavior and appetite. No mydriasis was noted in either eye at 24 hours and the anterior segment was normal. CONCLUSIONS: A combination of topical ophthalmic rocuronium bromide and 10% phenylephrine is safe and effective for mydriasis in juvenile loggerhead turtles.

Diverse marine animals migrate across vast expanses of seemingly featureless ocean before returning as adults to reproduce in the area where they originated. How animals accomplish such feats of natal homing is an enduring mystery. Growing evidence suggests, however, that sea turtles and salmon imprint on the magnetic field of their home area when young and then use this information to return as adults. Both turtles and salmon have the sensory abilities needed to detect the unique ‘magnetic signature’ of a coastal area. Analyses have revealed that, for both groups of animals, subtle changes in the geomagnetic field of the home region are correlated with changes in natal homing behavior. In turtles, a relationship between population genetic structure and the magnetic fields that exist at nesting beaches has also been detected, consistent with the hypothesis that turtles recognize their natal areas on the basis of magnetic cues. Salmon likely use a biphasic navigational strategy in which magnetic cues guide fish through the open sea and into the proximity of the home river where chemical cues allow completion of the spawning migration. Similarly, turtles may also exploit local cues to help pinpoint nesting areas once they have arrived in the vicinity. Throughout most of the natal homing migration, however, magnetic navigation appears to be the primary mode of long-distance guidance in both sea turtles and salmon.

The scientific exploration of whether and how migratory animals return to their birth areas goes back at least to John James Audubon, who tied silver threads to the legs of young songbirds and obse...

In the beginning there was great confusion about animal migration. Aristotle, noting that the types of birds around him changed with the seasons, concluded that summer redstarts turned into robins at the onset of winter, and that garden warblers became blackcaps [1]. Others thought that birds disappear in winter because they hibernate submerged in mud. In a case of art decidedly not imitating life, a 16th century illustration accompanying the writings of Swedish Archbishop Olaus Magnus showed a fishing net filled with hibernating swallows being pulled from a lake [1]. Gradually, over centuries, these fanciful early explanations gave way to an understanding that migration is a widespread phenomenon and that Earth is alive with itinerant animals traversing continents, seas, and skies (Figure 1).

The San Cristóbal lava lizard, Microlophus bivittatus, is one of nine species of lava lizards endemic to the Galápagos Islands of Ecuador. No information presently exists about baseline health parameters for any of these species. We analysed blood samples drawn from 47 lizards (25 males and 22 females) captured at two locations on San Cristóbal Island. A portable blood analyser (iSTAT) was used to obtain near-immediate field results for total CO2, lactate, sodium, potassium, ionized calcium, glucose and haemoglobin. Standard laboratory haematology techniques were employed for differential white blood cell counts and haematocrit determination. Body temperature, heart rate and body measurements were also recorded. We found significant differences in haematocrit values between males and females. The values reported in this study provide baseline data that may be useful in detecting changes in health status among lava lizards affected by natural disturbances or anthropogenic threats. Our findings might also be helpful in future efforts to demonstrate associations between specific biochemical or haematological parameters and disease. Because there are several related species on different islands in the Galápagos archipelago, comparisons between populations and species will be of interest. Lay Summary:Haematology and biochemistry values of the San Cristóbal lava lizard Microlophus bivittatus, along with several other health parameters (morphometrics and temperature), are reported for the first time.

A correction to this article has been published and is linked from the HTML and PDF versions of this paper. The error has been fixed in the paper.

Diverse animals can orient to the earth's magnetic field (1-6), but the mechanism or mechanisms undrlying magnetic field detection have not been determined. Behavioral (7-9) amd neurophysiological (10-12) results suggest that the transduction process underlying magnetic compass orientation in vertebrates is light-dependent, a finding consistent with theoretical models proposing that magnetoreception involves a modulation of the response of retinal photoreceptors to light (13, 14). We report, however, that leatherback sea turtle (Dermochelys coriacea) hatchlings orient to the geomagnetic field in complete darkness. Thus, light-dependence is not a universal feature of vertebrate magnetic compasses.

Physiology as Integrated Systems Amanda Southwood Williard Vision Kerstin A. Fritsches and Eric J. Warrant Natal Homing and Imprinting in Sea Turtles Kenneth J. Lohmann, Catherine M.F. Lohmann, J. Roger Brothers, and Nathan F. Putman The Skeleton: An In Vivo View of Structure Jeanette Wyneken Age and Age Estimation in Sea Turtles Larisa Avens and Melissa L. Snover Molecular Genetics of Sea Turtles Michael P. Jensen, Nancy N. FitzSimmons, and Peter H. Dutton Oceanic Habits and Habitats: Dermochelys coriacea Vincent S. Saba Oceanic Habits and Habitats: Caretta caretta Katherine L. Mansfield and Nathan F. Putman Feeding Biology: Advances from Field-Based Observations, Physiological Studies, and Molecular Techniques T. Todd Jones and Jeffrey A. Seminoff Predators, Prey, and the Ecological Roles of Sea Turtles Michael R. Heithaus Exposure to and Effects of Persistent Organic Pollutants Jennifer M. Keller Fisheries Bycatch of Marine Turtles: Lessons Learned from Decades of Research and Conservation Rebecca Lewison, Bryan Wallace, Joana Alfaro-Shigueto, Jeff Mangel, Sara Maxwell, and Elliott Hazen Climate Change and Marine Turtles Mark Hamann, Mariana M.P.B. Fuentes, Natalie C. Ban, and Veronique J.L. Mocellin Free-Ranging Sea Turtle Health Mark Flint Sea Turtle Epibiosis Michael G. Frick and Joseph B. Pfaller Parasites of Marine Turtles Ellis C. Greiner Index

Organisms use a variety of environmental cues to orient their movements in three-dimensional space. Here, we show that the upward movement of young Chinook salmon ( Oncorhynchus tshawytscha ) emerging from gravel nests is influenced by the geomagnetic field. Fish in the ambient geomagnetic field travelled farther upwards through substrate than did fish tested in a field with the vertical component inverted. This suggests that the magnetic field is one of several factors that influences emergence from the gravel, possibly by serving as an orientation cue that helps fish determine which way is up. Moreover, our work indicates that the Oncorhynchus species are sensitive to the magnetic field throughout their life cycles, and that it guides their movements across a range of spatial scales and habitats.

During long-distance migrations, animals navigate using a variety of sensory cues, mechanisms and strategies. Although guidance mechanisms are usually studied under controlled laboratory conditions, such methods seldom allow for navigation behavior to be examined in an environmental context. Similarly, although realistic environmental models are often used to investigate the ecological implications of animal movement, explicit consideration of navigation mechanisms in such models is rare. Here, we used an interdisciplinary approach in which we first conducted lab-based experiments to determine how hatchling loggerhead sea turtles (Caretta caretta) respond to magnetic fields that exist at five widely separated locations along their migratory route, and then studied the consequences of the observed behavior by simulating it within an ocean circulation model. Magnetic fields associated with two geographic regions that pose risks to young turtles (due to cold wintertime temperatures or potential displacement from the migratory route) elicited oriented swimming, whereas fields from three locations where surface currents and temperature pose no such risk did not. Additionally, at locations with fields that elicited oriented swimming, simulations indicate that the observed behavior greatly increases the likelihood of turtles advancing along the migratory pathway. Our findings suggest that the magnetic navigation behavior of sea turtles is intimately tied to their oceanic ecology and is shaped by a complex interplay between ocean circulation and geomagnetic dynamics.

SummaryNatal homing is a pattern of behavior in which animals migrate away from their geographic area of origin and then return to reproduce in the same location where they began life [1–3]. Although diverse long-distance migrants accomplish natal homing [1–8], little is known about how they do so. The enigma is epitomized by loggerhead sea turtles (Caretta caretta), which leave their home beaches as hatchlings and migrate across entire ocean basins before returning to nest in the same coastal area where they originated [9, 10]. One hypothesis is that turtles imprint on the unique geomagnetic signature of their natal area and use this information to return [1]. Because Earth’s field changes over time, geomagnetic imprinting should cause turtles to change their nesting locations as magnetic signatures drift slightly along coastlines. To investigate, we analyzed a 19-year database of loggerhead nesting sites in the largest sea turtle rookery in North America. Here we report a strong association between the spatial distribution of turtle nests and subtle changes in Earth’s magnetic field. Nesting density increased significantly in coastal areas where magnetic signatures of adjacent beach locations converged over time, whereas nesting density decreased in places where magnetic signatures diverged. These findings confirm central predictions of the geomagnetic imprinting hypothesis and provide strong evidence that such imprinting plays an important role in natal homing in sea turtles. The results give credence to initial reports of geomagnetic imprinting in salmon [11, 12] and suggest that similar mechanisms might underlie long-distance natal homing in diverse animals.

A protein complex found to align with the direction of a magnetic field could be a key piece in the puzzle of how animals detect magnetic fields.

Although sea turtles face significant pressure from human activities, some populations are recovering due to conservation programs, bans on the trade of turtle products, and reductions in bycatch. While these trends are encouraging, the status of many populations remains unknown and scientific monitoring is needed to inform conservation and management decisions. To address these gaps, this study presents methods for using unmanned aerial systems (UAS) to conduct population assessments. Using a fixed-wing UAS and a modified strip-transect method, we conducted aerial surveys along a three-kilometer track line at Ostional, Costa Rica during a mass-nesting event of olive ridley turtles (Lepidochelys olivacea). We visually assessed images collected during six transects for sea turtle presence, resulting in 682 certain detections. A cumulative total of 1091 certain and probable turtles were detected in the collected imagery. Using these data, we calculate estimates of sea turtle density (km−2) in nearshore waters. After adjusting for both availability and perception biases, we developed a low-end estimate of 1299 ± 458 and a high-end estimate of 2086 ± 803 turtles km−2. This pilot study illustrates how UAS can be used to conduct robust, safe, and cost-effective population assessments of sea turtle populations in coastal marine ecosystems.

Diverse taxa use Earth's magnetic field to aid both short- and long-distance navigation. Study of these behaviors has led to a variety of postulated sensory and processing mechanisms that remain unconfirmed. Although several models have been proposed to explain and understand these mechanisms' underpinnings, they have not necessarily connected a putative sensory signal to the nervous system. Using mathematical software simulation, hardware testing and the computational neuroscience tool of dynamic neural fields, the present work implements a previously developed conceptual model for processing magnetite-based magnetosensory data. Results show that the conceptual model, originally constructed to stimulate thought and generate insights into future physiological experiments, may provide a valid approach to encoding magnetic field information. Specifically, magnetoreceptors that are each individually capable of sensing directional information can, as a population, encode magnetic intensity and direction. The findings hold promise both as a biological magnetoreception concept and for generating engineering innovations in sensing and processing.

Diverse animals use Earth's magnetic field in orientation and navigation, but little is known about the molecular mechanisms that underlie magnetoreception. Recent studies have focused on two possibilities: (i) magnetite-based receptors; and (ii) biochemical reactions involving radical pairs. We used RNA sequencing to examine gene expression in the brain of rainbow trout (Oncorhynchus mykiss) after exposure to a magnetic pulse known to disrupt magnetic orientation behaviour. We identified 181 differentially expressed genes, including increased expression of six copies of the frim gene, which encodes a subunit of the universal iron-binding and trafficking protein ferritin. Functions linked to the oxidative effects of free iron (e.g. oxidoreductase activity, transition metal ion binding, mitochondrial oxidative phosphorylation) were also affected. These results are consistent with the hypothesis that a magnetic pulse alters or damages magnetite-based receptors and/or other iron-containing structures, which are subsequently repaired or replaced through processes involving ferritin. Additionally, some genes that function in the development and repair of photoreceptive structures (e.g. crggm3, purp, prl, gcip, crabp1 and pax6) were also differentially expressed, raising the possibility that a magnetic pulse might affect structures and processes unrelated to magnetite-based magnetoreceptors.

The hawksbill turtle, Eretmochelys imbricata, is a marine chelonian with a circum-global distribution, but the species is critically endangered and has nearly vanished from the eastern Pacific. Although reference blood parameter intervals have been published for many chelonian species and populations, including nesting Atlantic hawksbills, no such baseline biochemical and blood gas values have been reported for wild Pacific hawksbill turtles. Blood samples were drawn from eight hawksbill turtles captured in near shore foraging locations within the Galápagos archipelago over a period of four sequential years; three of these turtles were recaptured and sampled on multiple occasions. Of the eight sea turtles sampled, five were immature and of unknown sex, and the other three were females. A portable blood analyzer was used to obtain near immediate field results for a suite of blood gas and chemistry parameters. Values affected by temperature were corrected in two ways: (i) with standard formulas and (ii) with auto-corrections made by the portable analyzer. A bench top blood chemistry analyzer was used to measure a series of biochemistry parameters from plasma. Standard laboratory haematology techniques were employed for red and white blood cell counts and to determine haematocrit manually, which was compared to the haematocrit values generated by the portable analyzer. The values reported in this study provide reference data that may be useful in comparisons among populations and in detecting changes in health status among Galápagos sea turtles. The findings might also be helpful in future efforts to demonstrate associations between specific biochemical parameters and disease or environmental disasters.

The hawksbill turtle, Eretmochelys imbricata, is a marine chelonian with a circum-global distribution, but the species is critically endangered and has nearly vanished from the eastern Pacific. Although reference blood parameter intervals have been published for many chelonian species and populations, including nesting Atlantic hawksbills, no such baseline biochemical and blood gas values have been reported for wild Pacific hawksbill turtles. Blood samples were drawn from eight hawksbill turtles captured in near shore foraging locations within the Galápagos archipelago over a period of four sequential years; three of these turtles were recaptured and sampled on multiple occasions. Of the eight sea turtles sampled, five were immature and of unknown sex, and the other three were females. A portable blood analyzer was used to obtain near immediate field results for a suite of blood gas and chemistry parameters. Values affected by temperature were corrected in two ways: (i) with standard formulas and (ii) with auto-corrections made by the portable analyzer. A bench top blood chemistry analyzer was used to measure a series of biochemistry parameters from plasma. Standard laboratory haematology techniques were employed for red and white blood cell counts and to determine haematocrit manually, which was compared to the haematocrit values generated by the portable analyzer. The values reported in this study provide reference data that may be useful in comparisons among populations and in detecting changes in health status among Galápagos sea turtles. The findings might also be helpful in future efforts to demonstrate associations between specific biochemical parameters and disease or environmental disasters.

For sea turtles, an ability to detect land masses from a considerable distance away, and to distinguish coastal areas from the open sea, might be adaptive. The loggerhead turtle, Caretta caretta, can detect airborne odorants associated with food. To investigate whether sea turtles can also detect odors associated with land, we studied the responses of juvenile loggerheads to odors from coastal mud. Turtles were tested in a water-filled arena in which odorants could be introduced to the air above the water surface. Turtles exposed to air that had passed over a cup containing mud spent more time with their noses out of the water than did control turtles exposed to air that had passed over a cup containing distilled water. The results demonstrate for the first time that loggerheads can detect airborne odorants associated with land, an ability that might play a role in foraging, navigation, or both.

SummaryIn the final phase of their spawning migration, Pacific salmon use chemical cues to identify their home river, but how they navigate from the open ocean to the correct coastal area has remained enigmatic [1]. To test the hypothesis that salmon imprint on the magnetic field that exists where they first enter the sea and later seek the same field upon return [2–4], we analyzed a 56-year fisheries data set on Fraser River sockeye salmon, which must detour around Vancouver Island to approach the river through either a northern or southern passageway [5, 6]. We found that the proportion of salmon using each route was predicted by geomagnetic field drift: the more the field at a passage entrance diverged from the field at the river mouth, the fewer fish used the passage. We also found that more fish used the northern passage in years with warmer sea surface temperature (presumably because fish were constrained to more northern latitudes). Field drift accounted for 16% of the variation in migratory route used, temperature 22%, and the interaction between these variables 28%. These results provide the first empirical evidence of geomagnetic imprinting in any species and imply that forecasting salmon movements is possible using geomagnetic models.

Sea turtles are capable of navigating across large expanses of ocean to arrive at remote islands for nesting, but how they do so has remained enigmatic. An interesting example involves green turtles (Chelonia mydas) that nest on Ascension Island, a tiny land mass located approximately 2000 km from the turtles' foraging grounds along the coast of Brazil. Sensory cues that turtles are known to detect, and which might hypothetically be used to help locate Ascension Island, include the geomagnetic field, airborne odorants, and waterborne odorants. One possibility is that turtles use magnetic cues to arrive in the vicinity of the island, then use chemical cues to pinpoint its location. As a first step toward investigating this hypothesis, we used oceanic, atmospheric, and geomagnetic models to assess whether magnetic and chemical cues might plausibly be used by turtles to locate Ascension Island. Results suggest that waterborne and airborne odorants alone are insufficient to guide turtles from Brazil to Ascension, but might permit localization of the island once turtles arrive in its vicinity. By contrast, magnetic cues might lead turtles into the vicinity of the island, but would not typically permit its localization because the field shifts gradually over time. Simulations reveal, however, that the sequential use of magnetic and chemical cues can potentially provide a robust navigational strategy for locating Ascension Island. Specifically, one strategy that appears viable is following a magnetic isoline into the vicinity of Ascension Island until an odor plume emanating from the island is encountered, after which turtles might either: (1) initiate a search strategy; or (2) follow the plume to its island source. These findings are consistent with the hypothesis that sea turtles, and perhaps other marine animals, use a multi-modal navigational strategy for locating remote islands.

The Caribbean spiny lobster, Panulirus argus, is a migratory crustacean that uses Earth's magnetic field as a navigational cue, but how lobsters detect magnetic fields is not known. Magnetic material thought to be magnetite has previously been detected in spiny lobsters, but its role in magnetoreception, if any, remains unclear. As a first step toward investigating whether lobsters might have magnetite-based magnetoreceptors, we subjected lobsters to strong, pulsed magnetic fields capable of reversing the magnetic dipole moment of biogenic magnetite crystals. Lobsters were subjected to a single pulse directed from posterior to anterior and either: (1) parallel to the horizontal component of the geomagnetic field (i.e., toward magnetic north); or (2) antiparallel to the horizontal field (i.e., toward magnetic south). An additional control group was handled but not subjected to a magnetic pulse. After treatment, each lobster was tethered in a water-filled arena located within 200 m of the capture location and allowed to walk in any direction. Control lobsters walked in seemingly random directions and were not significantly oriented as a group. In contrast, the two groups exposed to pulsed fields were significantly oriented in approximately opposite directions. Lobsters subjected to a magnetic pulse applied parallel to the geomagnetic horizontal component walked westward; those subjected to a pulse directed antiparallel to the geomagnetic horizontal component oriented approximately northeast. The finding that a magnetic pulse alters subsequent orientation behavior is consistent with the hypothesis that magnetoreception in spiny lobsters is based at least partly on magnetite-based magnetoreceptors.

We review how ocean currents are measured (in both Eulerian and Lagrangian frameworks), how they are inferred from satellite observations, and how they are simulated in ocean general circulation models (OGCMs). We then consider the value of these 'direct' (in situ) and 'indirect' (inferred, simulated) approaches to biologists investigating current-induced drift of strong-swimming vertebrates as well as dispersion of small organisms in the open ocean. We sub- sequently describe 2 case studies. In the first, OGCM-simulated currents were compared with satellite-derived currents; analyses suggest that the 2 methods yield similar results, but that each has its own limitations and associated uncertainty. In the second analysis, numerical methods were tested using Lagrangian drifter buoys. Results indicated that currents simulated in OGCMs do not capture all details of buoy trajectories, but do successfully resolve most general aspects of current flows. We thus recommend that the errors and uncertainties in ocean current measure- ments, as well as limitations in spatial and temporal resolution of the surface current data, need to be considered in tracking studies that incorporate oceanographic data. Whenever possible, cross- validation of the different methods (e.g. indirect estimates versus buoy trajectories) should be undertaken before a decision is reached about which technique is best for a specific application.

In numerous studies involving hatchling sea turtles, researchers have collected small numbers of hatchlings from nests a few hours before the turtles would otherwise have emerged naturally. This procedure makes it possible to do experiments in which the behavioral or physio- logical responses of numerous hatchlings must be tested in a limited period of time, and also allows hatchlings to be released back into the sea in time to migrate offshore before dawn. In prin- ciple, however, the procedure might inadvertently reduce nest productivity (the number of hatch- lings that successfully leave the nest), if digging into a nest prior to emergence somehow reduces the ability of the remaining turtles to emerge. We compared nest productivity in 67 experimental loggerhead nests, from which we removed 10 hatchlings before a natural emergence, to 95 control nests left undisturbed before a natural emergence. The 2 groups showed no statistical differences in productivity. We conclude that taking a few hatchlings from a loggerhead nest shortly before a natural emergence has no negative impact on hatchling production if sampling is done with care at locations where there are few nest predators, and at sites where an emergence can be predicted because nest deposition dates are known.

Adults of the nudibranch mollusc Tritonia diomedea crawl using mucociliary locomotion. Crawling is controlled in part by the large Pedal 5 (Pd5) and Pedal 6 (Pd6) neurons that produce TritoniaPedal peptides (TPeps). TPeps elicit an increase in ciliary beat frequency, thereby increasing crawling speed. In adults of T. diomedea, an extensive network of TPep-containing neurites adjacent to the basement membrane of the pedal epithelium delivers TPeps to the ciliated cells. In this study, we show that diverse nudibranchs all have a pattern of TPep-like immunoreactivity similar to that of T. diomedea, with thin tracts of TPep-like immunoreactive (TPep-LIR) neurites projecting to the epithelial layer. We also show that members of two non-nudibranch gastropod species have a pattern of TPep-innervation similar to that of the nudibranchs. In addition, we characterized two pairs of motor neurons in adults of the nudibranch Armina californica that are possible homologues of the Pd5 and Pd6 cells in T. diomedea. Activity in one of these pairs, the Pedal Peptidergic Dorsal 1 (PPD1) cells, was correlated with mucociliary locomotion. The second pair, the Pedal Peptidergic Ventral 1 cells, shared synchronous synaptic input with the PPD1 cells, a pattern consistent with the shared synaptic input of the T. diomedea Pd5 and Pd6 cells. These findings suggest that the roles of the Pd5 and Pd6 cells as mucociliary motor neurons in nudibranchs are conserved evolutionarily. Additionally, the extensive network of TPep-LIR neurites seen in the foot of T. diomedea appears likely to be a common feature among gastropods.

SUMMARY Young loggerhead sea turtles (Caretta caretta) from the east coast of Florida, USA, undertake a transoceanic migration around the North Atlantic Gyre, the circular current system that flows around the Sargasso Sea. Previous experiments indicated that loggerhead hatchlings, when exposed to magnetic fields replicating those that exist at five widely separated locations along the migratory pathway, responded by swimming in directions that would, in each case, help turtles remain in the gyre and advance along the migratory route. In this study, hatchlings were exposed to several additional magnetic fields that exist along or outside of the gyre's northern boundary. Hatchlings responded to fields that exist within the gyre currents by swimming in directions consistent with their migratory route at each location, whereas turtles exposed to a field that exists north of the gyre had an orientation that was statistically indistinguishable from random. These results are consistent with the hypothesis that loggerhead turtles entering the sea for the first time possess a navigational system in which a series of regional magnetic fields sequentially trigger orientation responses that help steer turtles along the migratory route. By contrast, hatchlings may fail to respond to fields that exist in locations beyond the turtles' normal geographic range.