The Seasonal Aggregation of Whale Sharks at Ningaloo Reef, Western Australia: Currents, Migrations and the El Niño/ Southern Oscillation

Whale sharks, Rhincodon typus, seasonally aggregate in coastal waters off Ningaloo Reef, Western Australia. We review the oceanographic setting of the region and present evidence that such aggregations form as a result of migratory behavior associated with climatic and oceanographic processes. We utilise records of whale shark abundance collected at Ningaloo Reef from dedicated searches by boat and aircraft and from log sheets recorded by the tourism industry. Measures of whale shark abundance derived from log sheet data sets were moderately correlated with the Southern Oscillation Index and weakly correlated with coastal sea level, an index of the strength of the Leeuwin Current, and sea surface temperature over the period 1993 to 1998. Abundances of whale sharks derived from boat searches from 1983 to 1992 were also correlated with fluctuations in the Southern Oscillation Index, except during a three year period from 1988 to 1990. We conclude that, at least in some years, there appears to be a link between the abundance of aggregating whale sharks and the physical and biological oceanography of the region, with greater whale shark numbers in La Niña years. The lack of correlation in other years may be due to a combination of uneven quality of data and/or aggregations occurring in response to a complex interaction between the physical and biological oceanography of the region.     

Short‐term survival and movements of Atlantic sharpnose sharks captured by hook‐and‐line in the north‐east Gulf of Mexico

Ultrasonic telemetry was used to compare post‐release survival and movements of Atlantic sharpnose sharks Rhizoprionodon terraenovae in a coastal area of the north‐east Gulf of Mexico. Ten fish were caught with standardized hook‐and‐line gear during June to October 1999. Atlantic sharpnose sharks were continuously tracked after release for periods of 0·75 to 5·90 h and their positions recorded at a median interval of 9 min. Individual rate of movement was the mean of all distance and time measurements for each fish. Mean ± s.e . individual rate of movement was 0·45 ± 0·06 total lengths per second (L_T s^?1) and ranged from 0·28 to 0·92 L_T s^?1 over all fish. Movement patterns did not differ between jaw and internally hooked Atlantic sharpnose sharks. Individual rate of movement was inversely correlated with bottom water temperature at capture (r² = 0·52, P ≤ 0·05). No consistent direction in movement was detected for Atlantic sharpnose sharks after release, except that they avoided movement towards shallower areas. Capture‐release survival was high (90%), with only one fish not surviving, i.e. this particular fish stopped movement for a period of 10 min. Total rate of movement was total distance over total time (m min^?1) for each Atlantic sharpnose shark. Mean total rate of movement was significantly higher immediately after release at 21·5 m min^?1 over the first 1·5 h of tracking, then decreased to 11·2 m min^?1 over 1·5–6 h, and 7·7 m min^?1 over 3–6 h (P ≤ 0·002), which suggested initial post‐release stress but quick recovery from capture. Thus, high survival (90%) and quick recovery indicate that the practice of catch‐and‐release would be a viable method to reduce capture mortality for R. terraenovae.     

Do they stay or do they go? Acoustic monitoring of whale sharks at Ningaloo Marine Park,Western Australia

Whale sharks Rhincodon typus were monitored via acoustic transmitters at the northern end of Western Australia's Ningaloo Marine Park to establish the extent to which the species inhabits the region beyond the whale‐shark ecotourism industry season, which usually extends from March to August in each year. Despite the vast majority (c. 98%) of photographic submissions of R. typus from Ningaloo Reef being between March and August, acoustic detections from the tagged R. typus at Ningaloo were recorded in all months of the year, but do not preclude the occurrence of extended absences. It is concluded that as a species, R. typus occurs year round at Ningaloo, where it generally remains in close proximity to the reef edge, but that some individuals move outside of the detection range of the array for extended periods.     

To go or not to go with the flow: Environmental influences on whale shark movement patterns

Seven whale sharks were tracked using satellite-linked tags from Ningaloo Reef, off northern Western Australia, following tagging in April and June 2002 and April-May 2005. We investigated how the movements of those whale shark tracks were influenced by geostrophic surface currents during sequential one-week periods by using a passive diffusion model parameterised with observed starting locations of the sharks and weekly maps of surface current velocity and direction (derived from altimetry). We compared the outputs from the passive diffusion model and maps of chlorophyll-a concentration (SeaWiFs/MODIS) and with the actual tracks of the sharks using GIS and generalized linear mixed-effects models (GLMM). The GLMM indicated very little support for passive diffusion with sea-surface ocean currents influencing whale shark distributions in the north eastern Indian Ocean. Moreover, the sharks' movements correlated only weakly with the spatial distribution of sea-surface chlorophyll-a concentrations. The seven whale sharks had average swimming speeds comparable with those recorded in other satellite tracking studies of this species. Swimming speeds of the seven sharks were similar to those reported in previous studies and up to three times greater than the maximum sea-surface current velocities that the sharks encountered while traversing into lower southerly latitudes (moving northward towards the equator). Our results indicate that whale sharks departing from Ningaloo travel actively and independently of near-surface currents where they spend most of their time despite additional metabolic costs of this behaviour.     

Genetic structure of populations of whale sharks among ocean basins and evidence for their historic rise and recent decline

This study presents genetic evidence that whale sharks, Rhincodon typus, are comprised of at least two populations that rarely mix and is the first to document a population expansion. Relatively high genetic structure is found when comparing sharks from the Gulf of Mexico with sharks from the Indo‐Pacific. If mixing occurs between the Indian and Atlantic Oceans, it is not sufficient to counter genetic drift. This suggests whale sharks are not all part of a single global metapopulation. The significant population expansion we found was indicated by both microsatellite and mitochondrial DNA. The expansion may have happened during the Holocene, when tropical species could expand their range due to sea‐level rise, eliminating dispersal barriers and increasing plankton productivity. However, the historic trend of population increase may have reversed recently. Declines in genetic diversity are found for 6 consecutive years at Ningaloo Reef in Australia. The declines in genetic diversity being seen now in Australia may be due to commercial‐scale harvesting of whale sharks and collision with boats in past decades in other countries in the Indo‐Pacific. The study findings have implications for models of population connectivity for whale sharks and advocate for continued focus on effective protection of the world's largest fish at multiple spatial scales.     

Bioenergetics of juvenile whitespotted bamboo shark Chiloscyllium plagiosum [Anonymous (Bennett)]

This study establishes the bioenergetics budget of juvenile whitespotted bamboo shark Chiloscyllium plagiosum by estimating the standard metabolic rate (R_S), measuring the effect of body size and temperature on the R_S, and identifying the specific dynamic action (R_SDA) magnitude and duration of that action in juvenile whitespotted bamboo sharks. The mean ±s .d . (R_S) of six fish (500–620 g) measured in a circular closed respirometry system was 30·21 ± 5·68 mg O₂ kg^?1 h^?1 at 18° C and 70·38 ± 14·81 mg O₂ kg^?1 h^?1 at 28° C, respectively. There were no significant differences in R_S between day and night at either 18 or 28° C (t‐test, P > 0·05). The mean ±s .d . Q₁₀ for 18–28° C was 2·32 ± 0·06 (n = 6). The amount of oxygen consumed per hour changed predictably with body mass (M; 295–750 g) following the relationship: (n = 40, r²= 0·92, P < 0·05). The mean magnitude of R_SDA was 95·28 ± 17·55 mg O₂ kg^?1 h^?1. The amount of gross ingested energy (E_I) expended as R_SDA ranged from 6·32 to 12·78% with a mean ±s .d . of 8·01 ± 0·03%. The duration of the R_SDA effect was 122 h. The energy content of juvenile whitespotted bamboo shark, squid and faeces determined by bomb calorimeter were 19·51, 20·3 and 18·62 kJ g dry mass^?1. A mean bioenergetic budget for juvenile whitespotted bamboo sharks fed with squid at 18° C was 100C = 29·5G + 31·9R_S+ 28·2R_SDA+ 6·7F + 2·1E + 1·6U, where C = consumption, G = growth, F = egestion, E = excretion and U = unaccounted energy.     

Oceanographic and atmospheric phenomena influence the abundance of whale sharks at Ningaloo Reef, Western Australia

Seasonal observations of whale shark abundance recorded by ecotourist operators at Ningaloo Reef, Western Australia from 1999 to 2004 were compared with weekly regional and global oceanographic and atmospheric variables, including average sea surface temperatures, along-shelf wind shear, sea level and the Southern Oscillation Index (SOI). Estimates of these physical variables were derived from either ground-based data or from remote sensing instruments. A generalised linear mixed-effects modelling (GLMM) approach with random sampling and model simulation was used to determine the relationships between the number of whale sharks and all model variants of the environmental parameters, using information-theoretic weights of evidence to rank models. SOI and wind shear had the most support for explaining the deviance in weekly whale shark abundance at Ningaloo Reef during a season. The SOI and wind shear variables positively influenced whale shark abundance such that more sharks were sighted when the Southern Oscillation was stronger and along-shelf winds were increasingly prevalent. This may reflect changes in the strength of oceanographic processes such as the Leeuwin Current (in response to the Southern Oscillation) and wind/current driven upwelling which may affect the abundance of whale sharks transported to the region and/or the availability of their prey by driving productivity changes.     

Habitat suitability of Rhincodon typus in three localities of the Gulf of California: Environmental drivers of seasonal aggregations

The whale shark is an endangered species that usually feeds in coastal areas of highly productive seas such as the Gulf of California, Mexico. This study aims to describe the effect of sea surface temperature, chlorophyll a, bathymetry and slope on the habitat suitability of whale sharks in three important aggregation sites of the Gulf of California. A total of 2396 records of occurrence of whale sharks were obtained from international databases and scientific literature between 1996 and 2018. These records were used for the creation of a species distribution model using MaxEnt for each of the three aggregation sites. The concentration of chlorophyll a explained 71% of the habitat suitability, followed by bathymetry and slope with a combined 17%, and sea surface temperature constituting 10% of the model. Habitat suitability was related to areas where nontargeted fisheries may impact whale sharks through bycatch, entanglement and ship strikes. The implications for the conservation of whale sharks should be considered for management decisions in terms of marine protected areas, fishing refugees or bans, and other regulations regarding fisheries activities.     

Multi-Year Impacts of Ecotourism on Whale Shark (Rhincodon typus) Visitation at Ningaloo Reef,Western Australia

In-water viewing of sharks by tourists has become a popular and lucrative industry. There is some concern that interactions with tourists with ecotourism operations might harm sharks through disruption of behaviours. Here, we analysed five years of whale shark (Rhincodon typus) encounter data by an ecotourism industry at Ningaloo Reef, Western Australia, to assess the impact of ecotourism interactions on shark visitation, within the context of the biological and physical oceanography of the region. Our data base consisted of 2823 encounter records for 951 individual whale sharks collected by ecotourism operators between 2007 and 2011. We found that total encounters per whale shark and encounters per boat trip increased through time. On average, whale sharks re-encountered in subsequent years were encountered earlier, stayed longer and tended to be encountered more often within a season than sharks that were only encountered in a single year. Sequential comparisons between years did not show any patterns consistent with disturbance and the rate of departure of whale sharks from the aggregation was negatively correlated to the number of operator trips. Overall, our analysis of this multi-year data base found no evidence that interactions with tourists affected the likelihood of whale shark re-encounters and that instead, physical and biological environmental factors had a far greater influence on whale shark visitation rates. Our approach provides a template for assessing the effects of ecotourism interactions and environmental factors on the visitation patterns of marine megafauna over multiple years.     

Killer whale abundance and predicted narwhal consumption in the Canadian Arctic

Range expansions and increases in the frequency of killer whale (Orcinus orca) sightings have been documented in the eastern Canadian Arctic, presumably the result of climate change‐related sea‐ice declines. However, the effects of increased predator occurrence on this marine ecosystem remain largely unknown. We explore the consequences of climate change‐related range expansions by a top predator by estimating killer whale abundance and their possible consumptive effects on narwhal (Monodon monoceros) in the Canadian Arctic. Individual killer whales can be identified using characteristics such as acquired scars and variation in the shape and size of their dorsal fins. Capture–mark–recapture analysis of 63 individually identifiable killer whales photographed between 2009 and 2018 suggests a population size of 163 ± 27. This number of killer whales could consume >1,000 narwhal during their seasonal residency in Arctic waters. The effects of such mortality at the ecosystem level are uncertain, but trophic cascades caused by top predators, including killer whales, have been documented elsewhere. These findings illustrate the magnitude of ecosystem‐level modifications that can occur with climate change‐related shifts in predator distributions.     

Foraging ecology of Cookiecutter Sharks (<Emphasis Type="Italic">Isistius brasiliensis</Emphasis>) on pelagic fishes in Hawaii,inferred from prey bite wounds

The Cookiecutter Shark (Isistius brasiliensis) is an ecto-parasitic predator of numerous large pelagic fish and mammals. However, little is known of its foraging ecology due to its elusive foraging tactics in the pelagic environment. We used bite scar patterns on pelagic fishes landed at the Honolulu Fish Auction to assess some of the Cookiecutter Shark foraging habits. Swordfish (Xiphias gladius) had the greatest percentage of bites (87.9 ± 25.0% of individuals had healed scars) followed by Opah (Lampris guttatus, 33.0 ± 8.3% of individuals). Most fish with scars only had one Cookiecutter Shark bite per individual with the exception of Swordfish, which often had >5 bites per individual. Furthermore, Swordfish had a higher proportion of healed bite scars meaning they had been attacked while free-swimming. Seasonal changes in the probability of hooked fish being bitten by sharks were apparent for Swordfish, Bigeye Tuna and Opah. Based on bite scar diameter, larger Cookiecutter Sharks may preferentially attack Swordfish rather than the other species of pelagic fish. When taken in conjunction with diving behavior of pelagic fish, and fishing depths, the results add further support to the hypothesis that Cookiecutter Sharks perform diel vertical migrations.     

A chemical modification of myeloperoxidase and lactoperoxidase with 2-(O-methoxypolyethylene glycol)-4,6-dichloro-s-triazine (activated PEG1)

The modification of myeloperoxidase and lactoperoxidase with 2-(O-methoxypolethylene glycol)-4, 6-dichloro-s-triazine, an activated polyethylene glycol (PEG₁), was investigated. The modification caused a shift of the Soret band in the light absorption spectrum, from 430 nm to 418 nm in the case of myeloperoxidase (native ferric form), and from 412 nm to 406 nm in the case of lactoperoxidase (native ferric form). PEG₁-modified myeloperoxidase and PEG₁-modified lactoperoxidase both failed to bind with antiserum to the respective native enzyme, but both retained respectively 4·5±0·3 per cent (mean±SE, n=5) and 0·6±0·2 per cent (mean±SE, n=5) of the activities of peroxidation of the hydrogen donor o-methoxyphenol in comparison with the native enzyme, and 1·5±0·2 per cent (mean±SE, n=5) and 1·2±0·2 per cent (mean±SE, n=5) of the activities of destruction of fuchsin basic in the presence of hydrogen peroxide and a halide, bromide. The pH dependencies of the peroxidating activities were almost the same as those of the corresponding native enzymes, but both the optimal pHs of the reactions involving the destruction of fuchsin basic were shifted by approximately 1·0 pH unit toward neutral pH compared with the respective native enzymes. © 1998 John Wiley & Sons, Ltd.     

Size frequency,dispersal distances and variable growth rates of young sharks in a multi-species aggregation

During a mark–recapture survey from November 2014 until April 2017, 333 neonatal and juvenile blacktip reef sharks Carcharhinus melanopterus and 302 neonatal and juvenile sicklefin lemon sharks Negaprion acutidens were tagged and measured at the uninhabited and isolated St. Joseph Atoll (Republic of Seychelles). Both species demonstrated seasonal reproductive synchronicity and relatively large sizes at birth. Despite the extended times at liberty > 2.5 years, the majority of recaptures were found in close proximity to the initial tagging location (< 500 m). Annual growth rates of C. melanopterus (n = 24) and N. acutidens (n = 62) ranged from 6.6 to 31.7 cm year⁻¹ (mean ± SE; 16.2 ± 1.2 cm year⁻¹) and 0.2 to 32.2 cm year⁻¹ (11.8 ± 1 cm year⁻¹), respectively and are to date the most variable ever recorded in wild juvenile sharks. High abundances of both species coupled with long-term and repeated recaptures are indicative of a habitat where juveniles can reside for their first years of life. However, large variability in annual growth rates in both species may suggest high intra and interspecific competition induced by a possibly resource limited, isolated habitat.     

Aggregations of juvenile whale sharks (<Emphasis Type="Italic">Rhincodon typus</Emphasis>) in the Gulf of Tadjoura,Djibouti

A total of 23 whale sharks were identified over a 5 d period in the Arta Bay region of the Gulf of Tadjora, Djibouti. Most of the sharks aggregating in this area were small (<4 m TL) males. Individuals were identified using photographs of distinctive scars and spot and stripe patterns on the sides of the animals. Of these, 65% had scarring that was attributable to boat or propeller strikes. Most of the whale sharks we encountered were feeding on dense accumulations of plankton in shallow water just off (10–200 m) the shoreline. This food source may account for the aggregation of sharks in this area. One 3 m male shark was tagged with an ARGOS (Splash) satellite tag for 9 d. During this time the shark traversed to the shoreline on the opposite side of the Gulf (a distance of 14 km) and then returned to the Arta Bay area before retracing his path to the other shore. The shark spent most of the daylight hours at the surface, while at night dives were more frequent, deeper and for longer durations.

Biochemical composition of a dominant detritivorous fish Prochilodus lineatus along pollution gradients in the Paraná-Río de la Plata Basin

The biochemical composition of muscle, liver and stomach contents of a detritivorous fish Prochilodus lineatus was analysed and compared to settling particles and sediments along pollution gradients over 1500 km of the Río de la Plata Basin to evaluate the effects of anthropogenic discharges in a detritus food chain. The stomach contents of P. lineatus collected in the polluted Metropolitan Buenos Aires coast were enriched in proteins, carbohydrates and lipids, similar to settling particulates collected in the sewer area, and two to five times higher than underlying sediments, supporting the interpretation that P. lineatus feeds on unconsolidated organic flocs freshly decanted from mixed industrial and sewage outfalls. Fish from Buenos Aires had consistently higher standard length (L_S) and mass (M_T) slopes (b = 3·5), condition indexes (K = 3·01 ± 0·47, mean ±s.d .) and muscle fat content (fat = 23·8 ± 13·8% wet mass, mean ±s.d .) relative to northern fish (b = 2·7, K = 2·22 ± 0·39, fat = 3·4 ± 3·2% wet mass, respectively), suggesting that sewage‐derived organic matter was an enriched diet, which allowed an enhanced body mass gain and fat accumulation compared to organic‐poor vegetal detritus in the north Paraná area. Buenos Aires fish also showed higher hepato‐somatic indices (mean ±s.d . I_H 1·41 ± 0·49 v. 0·70 ± 0·32, respectively), which correlated with their two to three orders of magnitude higher hydrocarbon and polychlorinated biphenyl (PCB) loads, suggesting an enhanced detoxifying metabolism. The northward migration of fatty P. lineatus was evidenced by the presence of clear outliers in the L_S and M_T relationship, K and fat content along the Paraná River.     

Using visible implant elastomer to study ammocoete populations with Cormack–Jolly–Seber models

This study examined the efficacy of marking wild populations of lampreys with visible implant elastomer (VIE) for 6–18 months to examine ammocoete movements using Cormack–Jolly–Seber (CJS) open‐population models. These methods were tested on two lamprey populations in different river systems. American brook lamprey Lethenteron appendix at Dyke Creek apparent survival (φ) was high in the summer and winter (c. 0·7), but declined after flow events in the spring and autumn. Sea lamprey Petromyzon marinus at Oquaga Creek φ in the top‐ranked models varied with stream location and time. Estimates of φ were similar to Dyke Creek during the summer (c. 0·7), but declined after flow events and remained low (c. 0·1) in winter. Open‐population models support current understanding of ammocoete movement, i.e. dispersal is driven by high‐flow events at certain times of the year. The present study provides a framework to study ammocoetes with VIE.     

No overall benefit of predator exclosure cages for the endangered St. Helena Plover Charadrius sanctaehelenae

Predator exclosure cages are designed to increase the clutch survival of ground‐nesting birds. Predator exclosures provided for the endangered St. Helena Plover Charadrius sanctaehelenae, however, did not result in differences in clutch survival between protected and control nests and may have resulted in elevated adult mortality. Exclosures did not exclude all cats, the dominant nest predator, and it is likely that cats caused the adult mortalities observed close to the exclosures. A population model indicates that even if predator exclosures had excluded all cats, the benefits of increased clutch survival would have been more than negated by the estimated decrease in adult survival. The overall effect of predator exclosures needs to be clarified for other species, taking into consideration annual productivity and adult survival, to understand the circumstances in which predator exclosures are beneficial.     

Combined Effects of Virus, Pesticide, and Predator Cue on the Larval Tiger Salamander (Ambystoma tigrinum)

Emerging diseases and environmental contamination are two of the leading hypotheses for global amphibian declines. Yet few studies have examined the influence of contaminants on disease susceptibility, and even fewer have incorporated the role of natural stressors such as predation. We performed a factorial study investigating the interaction of the insecticide carbaryl, dragonfly predator cue, and the emerging pathogen Ambystoma tigrinum virus (ATV) on fitness correlates and disease susceptibility in tiger salamander larvae. Four week old larvae were exposed for 22 days in a 2 (0, 500 μg/l carbaryl) × 2 (control, predator cue water) × 2 (0, 1 × 10⁴ pfu ATV) factorial designed laboratory study. Results show significant impacts to survival of larvae for both virus and predator cue treatments, as well as an interactive effect between the two, in which predator cue strongly exacerbated disease-driven mortality. There was a clear pattern of reduced survival with the addition of stressors, with those where all three stressors were present exhibiting the worst effects (a decrease in survival from 93 to 60%). On those that survived, we also detected several sub-lethal impacts in mass, SVL, and development. Predator cue and pesticide treatments significantly reduced both SVL and mass. Virus and predator treatments significantly slowed development. Stressors also exhibited opposing effects on activity. Predator cue caused a significant reduction in activity, whereas virus caused a significant increase in activity over time. These results highlight the importance of examining combined natural and introduced stressors to understand potential impacts on amphibian species. Such stressors may contribute to the emergence of ATV in particular regions, raising concerns about the influence of pesticides on disease emergence in general.     

Freshwater influences on embryos,hatching and larval survival of euryhaline Gulf killifish Fundulus grandis and potential constraints on habitat distribution

The influence of fresh water on potential habitat occupancy of early life‐history stages of euryhaline Gulf killifish Fundulus grandis was determined by evaluating fertilization of freshwater‐spawned eggs and subsequent survival of embryos and larvae in comparison with saline water (salinity 7). Overall per cent fertilization of eggs was low (mean ± s.e . = 20·21 ± 0·03%). Embryo survival was greater in saline water, but hatching rate (mean ± s.e . = 81·6 ± 0·1%) and post‐hatch survival of larvae in fresh water (mean ± s.e . = 74·5 ± 0·1%) was relatively high. Therefore, the relative limitation of fresh water on habitat distribution of F. grandis changes with development, stimulating further questions on factors that may constrain habitat distribution of euryhaline fishes.