Ensemble modeling of the potential distribution of the whale shark in the Atlantic Ocean

The whale shark (Rhincodon typus) is an endangered marine fish species which can be adversely affected by the fishing activities of the industrial purse seine fleet targeting tropical tuna. Tuna tend to aggregate around all types of floating objects, including whale sharks. We analyzed and modeled the spatial distribution and environmental preferences of whale sharks based on the presence and absence data from fishing observations in the Atlantic Ocean. We used a thorough multialgorithm analysis, based on a new presence–absence dataset, and endeavored to follow the most recent recommendations on best practices in species distribution modeling. First, we selected a subset of relevant variables using a generalized linear model that addressed multicollinearity, statistical errors, and information criteria. We then used the selected variables to build a model ensemble including 19 different algorithms. After eliminating models with insufficient performance, we assessed the potential distribution of whale sharks using the mean of the predictions of the selected models. We also assessed the variance among the predictions of different algorithms, in order to identify areas with the highest model consensus. The results show that several coastal regions and warm shallow currents, such as the Gulf Stream and the Canary and Benguela currents, are the most suitable areas for whale sharks under current environmental conditions. Future environmental projections for the Atlantic Ocean suggest that some of the suitable regions will shift northward, but current concentration areas will continue to be suitable for whale shark, although with less productivity, which could have negative consequences for conservation of the species. We discuss the implications of these predictions for the conservation and management of this charismatic marine species.     

Feeding habits of the whale shark (<Emphasis Type="Italic">Rhincodon typus</Emphasis>) inferred by fatty acid profiles in the northern Mexican Caribbean

Whale shark (Rhincodon typus, Smith, 1828) is an endangered species with anthropogenic pressures due to increasing demand of encounter tourism activities. Research efforts to identify management and conservation strategies for this species are needed. The Northern Mexican Caribbean is one of the most important feeding aggregation sites of whale sharks worldwide. In this study, Mexican Caribbean whale shark feeding habits are assessed by means of fatty acid (FA) signature analysis, a biochemical non-destructive technique widely applied in trophic ecology studies. Sub-dermal tissue biopsies of 68 whale sharks and samples of their potential prey (zooplankton) were collected during 2010 and 2011 in two areas with high R. typus abundance. Zooplankton samples (n?=?17) were divided in two categories: mixed zooplankton (several groups of zooplankton) and fish eggs (> 95% of sample components were fish eggs). FA profiles of whale shark tissue sampled between years showed significant variability; while there was no intraspecific differences in FA signature related to sex, size and location. FA profiles of whale sharks and their potential prey were dominated by saturated fatty acids (SFA). R. typus FA signature was significantly different from that of mixed zooplankton; on the other hand, whale shark and fish egg FA profiles formed groups with overlapping values and registered high levels of oleic acid. PUFA average ω3/ ω6 ratio on whale shark FA profiles for both years was below 1. Arachidonic acid (ARA) percentage was higher in whale shark biopsies (13.2% in 2010, 6.8% in 2011) compared to values observed in fish eggs (2.0%) and mixed zooplankton (1.4%). Similarity between FA profiles of whale sharks and fish eggs, low levels of bacterial FA found in R. typus biopsies, as well as whale shark feeding behavior observations in the study area, suggest that R. typus is feeding mainly on surface zooplankton in Mexican Caribbean; however, elevated ARA percentages in whale shark samples may indicate that this species has complementary feeding sources, such as demersal zooplankton, which has been reported in other aggregation sites. Results obtained contribute to the knowledge of the whale shark trophic ecology in the area, but are inconclusive. Further studies are recommended to evaluate whale shark FA profiles from different tissues (muscle or blood); also, broader information is needed about zooplankton FA signature in the study area.     

Whale killers: Prevalence and ecological implications of killer whale predation on humpback whale calves off Western Australia

Reports of killer whales (Orcinus orca) preying on large whales have been relatively rare, and the ecological significance of these attacks is controversial. Here we report on numerous observations of killer whales preying on neonate humpback whales (Megaptera novaeangliae) off Western Australia (WA) based on reports we compiled and our own observations. Attacking killer whales included at least 19 individuals from three stable social groupings in a highly connected local population; 22 separate attacks with known outcomes resulted in at least 14 (64%) kills of humpback calves. We satellite‐tagged an adult female killer whale and followed her group on the water for 20.3 h over six separate days. During that time, they attacked eight humpback calves, and from the seven known outcomes, at least three calves (43%) were killed. Overall, our observations suggest that humpback calves are a predictable, plentiful, and readily taken prey source for killer whales and scavenging sharks off WA for at least 5 mo/yr. Humpback “escorts” vigorously assisted mothers in protecting their calves from attacking killer whales (and a white shark, Carcharodon carcharias). This expands the purported role of escorts in humpback whale social interactions, although it is not clear how this behavior is adaptive for the escorts.     

Donsol: an important reproductive habitat for the world's largest fish Rhincodon typus?

Little is known about the reproductive ecology of the whale shark Rhincodon typus. On 15 March 2020, a free-swimming neonate whale shark was found in the shallow, coastal waters of Donsol, Philippines. The total length of the male shark was 60 cm, falling within the species known size-at-birth. This is the third occurrence of neonatal whale sharks reported in the area, and coupled with the occurrence of juveniles and adults, and anecdotal mating and precopulatory behaviour reported herein, the importance of the site for this endangered species is highlighted.     

A review of the biology and ecology of the whale shark

The information available on the biology and ecology of the whale shark is reviewed, and is updated from material published since 1986. Research work carried out on the seasonal aggregation of whale sharks at the Ningaloo Reef in Western Australia is summarized. Future research studies on whale sharks in the Ningaloo Marine Park are discussed in the context of management of sustainable whale shark interaction tourism.     

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.     

Effects of species biology on the historical demography of sharks and their implications for likely consequences of contemporary climate change

Climate variation is an important factor shaping the demographic histories of many marine species, though impacts likely differ depending on species life history, habitat preferences and ecology. Investigating how species responded to historic climate fluctuations may provide critical insights into a species’ response to current climate change. Despite their ecological diversity, shark species share many similar life history characteristics and may be especially vulnerable to anthropogenic and climate impacts. We compared patterns of genetic variability, mismatch distributions and demographic reconstructions from coalescence approaches among temperate and tropical shark species with differing ecological characteristics, to investigate the effect of the past glaciation cycles on population abundance. Genetic diversity at two mitochondrial DNA regions (ND2 and control region) was assayed in four North Pacific species, Pacific spiny dogfish, Pacific sleeper sharks, salmon shark, and bluntnose sixgill shark. In addition, control region sequences acquired from GenBank for five shark species [tope shark (California/Australia), white shark (California), blacktip shark (eastern and western Gulf of Mexico), lemon shark (Bahamas), and whale shark] were analyzed. General patterns in genetic diversity, mismatch analyses and Bayesian skyline plots supported our hypothesis that species biology affected the impact of climate variation on demographic history. Consequently, our results suggest that effects of contemporary climate change on sharks may be to some degree predictable from species biology, distribution, habitat and the impact of past climate events.     

Whale Shark (Rhincodon typus) Seasonal Presence,Residence Time and Habitat Use at Darwin Island,Galapagos Marine Reserve

The life history of the whale shark (Rhincodon typus), including its reproductive ecology, still remains largely unknown. Here, we present results from the first whale shark population study around Darwin Island, Galapagos Marine Reserve. Following a diversified approach we characterized seasonal occurrence, population structure and size, and described habitat use of whale sharks based on fine scale movements around the island. Whale shark presence at Darwin Island was negatively correlated with Sea Surface Temperature (SST), with highest abundance corresponding to a cool season between July and December over six years of monitoring. From 2011 to 2013 we photo-identified 82 whale sharks ranging from 4 to 13.1 m Total Length (TL). Size distribution was bimodal, with a great majority (91.5%) of adult female individuals averaging 11.35 m±0.12 m (TL±SE), all but one showing signs of a potential pregnancy. Population dynamics models for apparently pregnant sharks estimated the presence of 3.76±0.90 (mean ± SE) sharks in the study area per day with an individual residence time of 2.09±0.51 (mean ± SE) days. Movement patterns analysis of four apparently pregnant individuals tracked with acoustic tags at Darwin Island revealed an intense use of Darwin''s Arch, where no feeding or specific behavior has been recorded, together with periodic excursions around the island''s vicinity. Sharks showed a preference for intermediate depths (20–30 m) with occasional dives mostly to mid-water, remaining the majority of their time at water temperatures between 24–25°C. All of our results point to Darwin Island as an important stopover in a migration, possibly with reproductive purposes, rather than an aggregation site. Current studies carried out in this area to investigate regional scale movement patterns may provide essential information about possible pupping grounds for this enigmatic species.     

Apex predatory sharks and crocodiles simultaneously scavenge a whale carcass

Scavenging is an important component to the overall ecology of consumers in virtually all ecosystems on Earth. Given the energetic benefits of foraging on these resource subsidies, opportunistic predators will adjust their behaviors accordingly to maximize access. One of the many consequences of large-scale scavenging opportunities is species interactions that are rarely observed in nature. Here we describe the first published record of predatory sharks (tiger sharks, Galeocerdo cuvier) and saltwater crocodiles (Crocodylus porosus) foraging together in space and time, as documented on a large whale carcass off Western Australia. We report on and discuss the behaviors of the sharks and crocodiles in the hope of shedding new light on the interactions between apex predators that are rarely seen together, but may overlap under specific contexts.     

Movement Patterns of Juvenile Whale Sharks Tagged at an Aggregation Site in the Red Sea

Conservation efforts aimed at the whale shark, Rhincodon typus, remain limited by a lack of basic information on most aspects of its ecology, including global population structure, population sizes and movement patterns. Here we report on the movements of 47 Red Sea whale sharks fitted with three types of satellite transmitting tags from 2009–2011. Most of these sharks were tagged at a single aggregation site near Al-Lith, on the central coast of the Saudi Arabian Red Sea. Individuals encountered at this site were all juveniles based on size estimates ranging from 2.5–7 m total length with a sex ratio of approximately 1∶1. All other known aggregation sites for juvenile whale sharks are dominated by males. Results from tagging efforts showed that most individuals remained in the southern Red Sea and that some sharks returned to the same location in subsequent years. Diving data were recorded by 37 tags, revealing frequent deep dives to at least 500 m and as deep as 1360 m. The unique temperature-depth profiles of the Red Sea confirmed that several whale sharks moved out of the Red Sea while tagged. The wide-ranging horizontal movements of these individuals highlight the need for multinational, cooperative efforts to conserve R. typus populations in the Red Sea and Indian Ocean.     

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.     

Genetic profiling reveals illegal international trade in fins of the great white shark, <Emphasis Type="Italic">Carcharodon carcharias</Emphasis>

Great white sharks are protected by national legislation in several countries, making this species the most widely protected elasmobranch in the world. Although the market demand for shark fins in general has continued to grow, the value and extent of utilization of white shark fins in trade has been controversial. We combine law enforcement with genetic profiling to demonstrate that illegal trade in fins of this species is occurring in the contemporary international market. Furthermore, we document the presence of fins from very young white sharks in the trade, suggesting a multiple-use market (food to trophies) exists for fins of this species. The presence of small fins in the trade contradicts the view that white shark fins have market value only as large display trophies, and not as food. Our findings indicate that effective conservation of protected shark species will require international management regimes that include monitoring of the shark fishery and trade on a species-specific basis.     

Giant-toothed white sharks and cetacean trophic interaction from the Pliocene Caribbean Paraguaná Formation

The role of the extinct giant-toothed white sharkCarcharodon megalodon (Agassiz) in the Caribbean Neogene is discussed based on new evidence of predation on cetaceans from the Lower Pliocene Paraguaná Formation in Venezuela. Large sharks have occupied the highest trophic level in the marine environment. However, based on the recovery of a giant white shark tooth piercing a cetacean lumbar vertebra, the predator-prey relationship is discussed under the hypothesis of initial shark attack, subsequent floating transport of the cetacean carcass, and scavenging prior to deposition on the bottom. The scarce Caribbean fossil records of both giant-toothed sharks and cetaceans suggest that these species were transients during the Pliocene on the Venezuelan coast.      

New insights into whale shark <Emphasis Type="Italic">Rhincodon typus</Emphasis> diet in Brazil: an observation of ram filter-feeding on crab larvae and analysis of stomach contents from the first stranding in Bahia state

We describe the first record of a whale shark, Rhincodon typus, feeding in Brazilian coastal waters, and the first stranding record in the state of Bahia, Northeast Brazil. In April 2008, an individual of R. typus was observed surface feeding on Dromiidae crab larvae in the continental shelf off the coast of Bahia, near a gas platform. Other fishes were observed foraging in association with the whale shark. We also document the first stranding of R. typus on the coast of Bahia in October 2013. Biometric data confirmed that the stranded whale shark was a juvenile. Stomach content analysis revealed the ingestion of Geryonidae crab larvae. Plastic debris were also found in the gastric lumen of the stranded juvenile whale shark, and we speculate that it could have been a contributing factor to the stranding, and subsequent death of the whale shark. Crab larvae were observed in both of our records and likely to consist as relevant prey items for R. typus in Brazilian continental shelf. Our study provided a contribution on the diet and feeding behaviour of whale sharks in tropical oligotrophic waters and highlights the risks of marine pollution for the species conservation.     

Ocean‐scale prediction of whale shark distribution

Aim Predicting distribution patterns of whale sharks (Rhincodon typus, Smith 1828) in the open ocean remains elusive owing to few pelagic records. We developed multivariate distribution models of seasonally variant whale shark distributions derived from tuna purse‐seine fishery data. We tested the hypotheses that whale sharks use a narrow temperature range, are more abundant in productive waters and select sites closer to continents than the open ocean. Location Indian Ocean. Methods We compared a 17‐year time series of observations of whale sharks associated with tuna purse‐seine sets with chlorophyll a concentration and sea surface temperature data extracted from satellite images. Different sets of pseudo‐absences based on random distributions, distance to shark locations and tuna catch were generated to account for spatiotemporal variation in sampling effort and probability of detection. We applied generalized linear, spatial mixed‐effects and Maximum Entropy models to predict seasonal variation in habitat suitability and produced maps of distribution. Results The saturated generalized linear models including bathymetric slope, depth, distance to shore, the quadratic of mean sea surface temperature, sea surface temperature variance and chlorophyll a had the highest relative statistical support, with the highest percent deviance explained when using random pseudo‐absences with fixed effect‐only models and the tuna pseudo‐absences with mixed‐effects models (e.g. 58% and 26% in autumn, respectively). Maximum Entropy results suggested that whale sharks responded mainly to variation in depth, chlorophyll a and temperature in all seasons. Bathymetric slope had only a minor influence on the presence. Main conclusions Whale shark habitat suitability in the Indian Ocean is mainly correlated with spatial variation in sea surface temperature. The relative influence of this predictor provides a basis for predicting habitat suitability in the open ocean, possibly giving insights into the migratory behaviour of the world’s largest fish. Our results also provide a baseline for temperature‐dependent predictions of distributional changes in the future.     

The use of machine learning to detect foraging behaviour in whale sharks: a new tool in conservation

In this study we present the first attempt at modelling the feeding behaviour of whale sharks using a machine learning analytical method. A total of eight sharks were monitored with tri-axial accelerometers and their foraging behaviours were visually observed. Our results highlight that the random forest model is a valid and robust approach to predict the feeding behaviour of the whale shark. In conclusion this novel approach exposes the practicality of this method to serve as a conservation tool and the capability it offers in monitoring potential disturbances of the species.     

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.     

Segregation and foraging ecology of whale sharks, Rhincodon typus, in the southwestern Gulf of California

Sharks segregate by sex and size, but few studies have attempted to explain such behaviors. To address this, we examined aggregations and the foraging ecology of whale sharks in Bahía de La Paz (BLP) with aerial and ship surveys and direct observation. Zooplankton abundance and composition, and hydrographic conditions were analyzed in relation to whale shark occurrence to explore underlying factors causing segregations. We observed large aggregations of juveniles (<9 m total length, TL) inshore, comprised by 60 % male individuals, and small aggregations of adults (>9 m TL) offshore, composed of 84 % females. Juvenile sharks were associated to turbid shallow waters in BLP, where they performed stationary and dynamic suction feeding on dense copepod swarms. Adults occurred in oceanic waters and fed by ram-filtering on diffuse patches of euphausiids, with no association to oceanographic conditions. Such segregation may be advantageous to juvenile R. typus utilizing shallow coastal waters to find abundant preferred prey needed for their fast growth rates. Our studies suggest that the main driving forces of whale shark segregation by sex and size in BLP may be diet preference for juveniles and habitat preference for adult sharks.     

Feeding of the megamouth shark (Pisces: Lamniformes: Megachasmidae) predicted by its hyoid arch: A biomechanical approach

Studies of the megamouth shark, one of three planktivorous sharks, can provide information about their evolutionary history. Megamouth shark feeding has never been observed in life animals, but two alternative hypotheses on biomechanics suggest either feeding, i.e., ram feeding or suction feeding. In this study, the second moment of area of the ceratohyal cartilages, which is an indicator of the flexural stiffness of the cartilages, is calculated for 21 species of ram‐ and suction‐feeding sharks using computed tomography. The results indicate that suction‐feeding sharks have ceratohyal cartilages with a larger second moment of area than ram‐feeding sharks. The result also indicates that the ram–suction index, which is an indicator of relative contribution of ram and suction behavior, is also correlated with the second moment of area of the ceratohyal. Considering that large bending stresses are expected to be applied to the ceratohyal cartilage during suction, the larger second moment of area of the ceratohyal of suction‐feeding sharks can be interpreted as an adaptation for suction feeding. Based on the small second moment of area of the ceratohyal cartilage of the megamouth shark, the feeding mode of the megamouth shark is considered to be ram feeding, similar to the planktivorous basking shark. From these results, an evolutionary scenario of feeding mechanics of three species of planktivorous sharks can be suggested. In this scenario, the planktivorous whale shark evolved ram feeding from a benthic suction‐feeding ancestor. Ram feeding in the planktivorous megamouth shark and the basking shark evolved from ram feeding swimming‐type ancestors and that both developed their unique filtering system to capture small‐sized prey. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.     

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.