A. Peter Klimley

Philopatry and migration of Pacific white sharks

Advances in electronic tagging and genetic research are making it possible to discern population structure for pelagic marine predators once thought to be panmictic. However, reconciling migration patterns and gene flow to define the resolution of discrete population management units remains a major challenge, and a vital conservation priority for threatened species such as oceanic sharks. Many such species have been flagged for international protection, yet effective population assessments and management actions are hindered by lack of knowledge about the geographical extent and size of distinct populations. Combining satellite tagging, passive acoustic monitoring and genetics, we reveal how eastern Pacific white sharks (Carcharodon carcharias) adhere to a highly predictable migratory cycle. Individuals persistently return to the same network of coastal hotspots following distant oceanic migrations and comprise a population genetically distinct from previously identified phylogenetic clades. We hypothesize that this strong homing behaviour has maintained the separation of a northeastern Pacific population following a historical introduction from Australia/New Zealand migrants during the Late Pleistocene. Concordance between contemporary movement and genetic divergence based on mitochondrial DNA demonstrates a demographically independent management unit not previously recognized. This populations fidelity to discrete and predictable locations offers clear population assessment, monitoring and management options.

Movements and Swimming Behavior of Three Species of Sharks in La Jolla Canyon, California

We tracked six individuals of three shark species, the shortfin mako, Isurus oxyrinchus, great white, Carcharodon carcharias, and blue, Prionace glauca, near the submarine canyon off La Jolla, southern California during the summers of 1995 and 1997. The duration of tracking ranged from 2 to 38 h per shark. The mode of travel differed in one respect among species. The rate of movement of the endothermic species, the mako and white shark, exceeded that of the ectothermic species, the blue shark. Similarities among species were more common. Firstly, individuals of all three species swam in a directional manner. Secondly, individuals constantly moved up and down in the water column, exhibiting oscillatory or yo-yo swimming. Thirdly, members of the three species swam at the surface for prolonged periods. Finally, the movements of the mako and white sharks were at times loosely associated with bottom topography. We discuss the various adaptive advantages that have been proposed for these behavioral patterns. Oscillatory swimming has been attributed to the following: (1) heating the body in the warm surface waters after swimming in cold, deep water, (2) alternating between two strata of water, one carrying chemical information as to its source, and deriving a direction to that stratums origin, (3) conserving energy by quickly propelling oneself upward with many tail beats and slowly gliding downward with few beats, and (4) descending to where magnetic gradients are steeper, more perceptible, and useful to guide migratory movements. At the surface, an individual would be able to swim in a straight line by using following features as a reference: (1) celestial bodies, (2) polarized light, or (3) the earths main dipole field. Furthermore, an individual would conserve energy because of the greater ease to maintaining a warm body in the heated surface waters.

Evidence for Geomagnetic Imprinting as a Homing Mechanism in Pacific Salmon

In 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. 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, 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. 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.

Estimating Survival and Migration Route Probabilities of Juvenile Chinook Salmon in the Sacramento–San Joaquin River Delta

Abstract Juvenile Chinook salmon Oncorhynchus tshawytscha emigrating from natal tributaries of the Sacramento River must negotiate the Sacramento–San Joaquin River Delta, a complex network of natural and man-made channels linking the Sacramento River with San Francisco Bay. Natural processes and water management actions affect the fractions of the population using the different migration routes through the delta and survival within those routes. However, estimating these demographic parameters is difficult using traditional mark–recapture techniques, which depend on the physical recapture of fish (e.g., coded wire tags). Thus, our goals were to (1) develop a mark–recapture model to explicitly estimate the survival and migration route probabilities for each of four migration routes through the delta, (2) link these route-specific probabilities to population-level survival, and (3) apply this model to the first available acoustic telemetry data of smolt migration through the delta. The point estimate of sur...

A first estimate of white shark, Carcharodon carcharias, abundance off Central California

The decline of sharks in the global oceans underscores the need for careful assessment and monitoring of remaining populations. The northeastern Pacific is the home range for a genetically distinct clade of white sharks (Carcharodon carcharias). Little is known about the conservation status of this demographically isolated population, concentrated seasonally at two discrete aggregation sites: Central California (CCA) and Guadalupe Island, Mexico. We used photo-identification of dorsal fins in a sequential Bayesian mark–recapture algorithm to estimate white shark abundance off CCA. We collected 321 photographs identifying 130 unique individuals, and estimated the abundance off CCA to be 219 mature and sub-adult individuals ((130, 275) 95% credible intervals), substantially smaller than populations of other large marine predators. Our methods can be readily expanded to estimate shark population abundance at other locations, and over time, to monitor the status, population trends and protection needs of these globally distributed predators.

Spatiotemporal Patterns of White Shark (Carcharodon carcharias) Predation at the South Farallon Islands, California

Spatial and temporal records of 146 predatory attacks by white sharks (Carcharodon carcharias) on four species of pinnipeds, one bird, and one human at the South Farallon Islands, Central California, from late Aug. to early Dec. 198689 are presented. During each 3.5-mo period, attacks were (1) unevenly distributed in bouts separated by hiatuses in predation, (2) paired temporally within the same day, (3) at similar times and locations on consecutive days, and (4) all during daylight hours. Predation was observed most often within 450 m of shore, with a decrease in attack frequency with increasing depth. Within this high-risk zone, predation was concentrated near coastal departure and entry points of pinnipeds, and the predatory attack positions formed linear patterns leading away from the island. Consecutive predatory attacks were often near each other, yet at times alternated between localities on either side of the island.

Marine Migration of North American Green Sturgeon

Abstract An understanding of the distribution of North American green sturgeon Acipenser medirostris in coastal waters is crucial to minimize impacts on this vulnerable species from various fisheries. To determine migratory patterns, we tagged 213 subadult and adult green sturgeon in spawning rivers and summer aggregation areas with uniquely coded ultrasonic pingers and observed their coastal movements with arrays of automated hydrophones deployed along the West Coast of North America from southeast Alaska to Monterey Bay, California. Green sturgeon exhibited an annual migration along the continental shelf from U.S. to Canadian waters in the fall and an apparent return migration in the spring. Peak migration rates exceeded 50 km/d during the springtime southward migration. Large numbers of green sturgeon were detected near Brooks Peninsula on northwest Vancouver Island, British Columbia, during May-June and October-November. A single fish was detected in southeast Alaska in December. This pattern of detec...

Why publish Animal Biotelemetry

“Biotelemetry” refers to the sending of biological measurements from a transmitter on an animal to a receiver. The information is transmitted either with an ultrasonic (kHz) or radio signal (MHz). The ultrasonic signal is propagated by a ceramic ring, or piezo-electric transducer (PZT), whereas the radio signal is propagated by a conductive wire, or antenna. Which mode of telemetry is chosen to remotely monitor the behavior and movements of animals depends on the transmission properties of the medium. Marine scientists have utilized ultrasonic telemetry because a signal composed of those frequencies propagates quickly with little attenuation in salt water, whereas radio signals are absorbed quickly in this medium. Scientists working in the freshwater and terrestrial environments have mainly utilized radio telemetry because the signals of those frequencies propagate far in freshwater and air. Telemetry has often been used to study large marine vertebrates because poor visibility has limited direct observation. Tagged marine fish were first followed by boat, while determining the direction of the subject with a directional hydrophone and estimating its position by situating the boat above it and recording a position. The first transmitters were beacons that enabled researchers to describe the daily offshore migrations of tunas off Hawaii [1]. Later tags were equipped with multiple sensors such as those used to record the depths and headings of scalloped hammerhead sharks, and the water temperatures and underwater illumination to understand their navigational abilities [2]. Additional sensors continue to be developed such as an accelerometer used to identify behaviors from records of an individual’s movement along three axes [3] and pH sensors to detect feeding by a change in pH in the stomach associated with digestion [4]. Shipboard tracking of animals is an arduous activity. The recent development of coded beacons and small automated receivers have made it possible to describe simultaneously the “homing” behavior of multiple individuals to

Residency of the scalloped hammerhead shark (Sphyrna lewini) at Malpelo Island and evidence of migration to other islands in the Eastern Tropical Pacific

Sixty nine hammerhead sharks, Sphyrna lewini, were tagged at Malpelo Island (Colombia) with ultrasonic transmitters during March 2006, 2007 and 2008, as part of a study to understand their residency at the island and their horizontal and vertical movements. Five sharks visited Cocos Island, 627 km distant from Malpelo. One of the sharks that appeared at Cocos Island also visited the Galapagos Islands, 710 km from Cocos, a month later. There is connectivity of Sphyrna lewini between Malpelo, Cocos and the Galapagos Islands, but the frequency of movements between the islands appears to be relatively low (<7% of the tagged sharks). The most common depth at which the sharks swam coincided with the thermocline (rs = 0.72, p < 0.01). The depth of the thermocline varied depending on the time of the year. Nocturnal detections of the sharks were more frequent during the cold season than during the warm season (W = 60, p < 0.01). We also found that hammerheads spent significantly more time on the up-current side of the island (Kruskal-Wallis = 31.1008; p < 0.01). This study contributes to the knowledge of hammerhead sharks not only in Malpelo Island but also at a regional level in the Eastern Tropical Pacific.

Migration of green sturgeon, Acipenser medirostris, in the Sacramento River

Adult green sturgeon, Acipenser medirostris, were collected in San Pablo Bay, California, and surgically implanted with ultrasonic acoustic tags from 2004 to 2006. An array of automated acoustic monitors was maintained in the Sacramento River to record movements of these fish. We presumed movements to known spawning areas (based on previous green sturgeon egg collections) or areas with potential spawning habitat (characterized by substrate, flow, and temperature criteria) represented a “spawning migration.” Three separate annual “spawning migrations” were recorded involving 15 individuals. The majority of the Sacramento River migrants entered the system in the months of March and April. Two different patterns of “spawning migration” and out-migration were observed. Six individuals potentially spawned, over-summered and moved out of the river with the first fall flow event. This is believed to be the common behavior of the green sturgeon. Alternatively, nine individuals promptly moved out of the Sacramento River before 1 September, and any known flow or temperature cue. Some green sturgeon appeared to be impeded on their upstream movement by the 15 May closure of the Red Bluff Diversion Dam, and at least five passed under the dam gates during downstream migration. A delay in the closure of the Red Bluff Diversion Dam would likely allow upstream passage of spawning green sturgeon, further, the potential mortality affects of downstream passage beneath the Red Bluff Diversion Dam should be assessed. Specific protection should be also given to the large aggregation of green sturgeon located in the reach of the Sacramento River adjacent to the Glen Colusa Irrigation District pumping facility.