Inferred global connectivity of whale shark Rhincodon typus populations

Ten years have passed since the last synopsis of whale shark Rhincodon typus biogeography. While a recent review of the species' biology and ecology summarized the vast data collected since then, it is clear that information on population geographic connectivity, migration and demography of R. typus is still limited and scattered. Understanding R. typus migratory behaviour is central to its conservation management considering the genetic evidence suggesting local aggregations are connected at the generational scale over entire ocean basins. By collating available data on sightings, tracked movements and distribution information, this review provides evidence for the hypothesis of broad‐scale connectivity among populations, and generates a model describing how the world's R. typus are part of a single, global meta‐population. Rhincodon typus occurrence timings and distribution patterns make possible a connection between several aggregation sites in the Indian Ocean. The present conceptual model and validating data lend support to the hypothesis that R. typus are able to move among the three largest ocean basins with a minimum total travelling time of around 2–4 years. The model provides a worldwide perspective of possible R. typus migration routes, and suggests a modified focus for additional research to test its predictions. The framework can be used to trim the hypotheses for R. typus movements and aggregation timings, thereby isolating possible mating and breeding areas that are currently unknown. This will assist endeavours to predict the longer‐term response of the species to ocean warming and changing patterns of human‐induced mortality.     

Fisheries, management and conservation of the whale shark Rhincodon typus in Taiwan

The Taiwanese government introduced a catch-and-report system for the whale shark Rhincodon typus in 2001, and in the following year limited the total allowable catch (TAC) to 80 individuals. This limit was reduced over the following years, reaching a low of 30 individuals in 2007. In November 2007, a comprehensive ban on R. typus fishing came into effect and protection measures were adopted for the species. This study measured and recorded the total length (L(T) ), body mass and sex of 810 R. typus, using fisheries and other released data for the period 1995-2008. The mean L(T) of individuals caught from June to October was smaller than that for individuals caught from November to May. Mean annual catches in set nets were higher in south-western Taiwan than in other regions of Taiwan. The sex ratio showed that males were caught more frequently than females in Taiwanese waters, indicating sex segregation. For the first time, the stock structure, conservation and management regulation of R. typus in the north-west Pacific Ocean have been reviewed. The results provide a useful resource for future assessment of the possible development of R. typus ecotourism in Taiwan.     

A review of the biology, fisheries and conservation of the whale shark Rhincodon typus

Although the whale shark Rhincodon typus is the largest extant fish, it was not described until 1828 and by 1986 there were only 320 records of this species. Since then, growth in tourism and marine recreation globally has lead to a significant increase in the number of sightings and several areas with annual occurrences have been identified, spurring a surge of research on the species. Simultaneously, there was a great expansion in targeted R. typus fisheries to supply the Asian restaurant trade, as well as a largely un-quantified by-catch of the species in purse-seine tuna fisheries. Currently R. typus is listed by the IUCN as vulnerable, due mainly to the effects of targeted fishing in two areas. Photo-identification has shown that R. typus form seasonal size and sex segregated feeding aggregations and that a large proportion of fish in these aggregations are philopatric in the broadest sense, tending to return to, or remain near, a particular site. Somewhat conversely, satellite tracking studies have shown that fish from these aggregations can migrate at ocean-basin scales and genetic studies have, to date, found little graphic differentiation globally. Conservation approaches are now informed by observational and environmental studies that have provided insight into the feeding habits of the species and its preferred habitats. Notwithstanding these advances, there remain notable gaps in the knowledge of this species particularly with respect to the life history of neonates and adults who are not found in the feeding aggregations.     

First record of a whale shark Rhincodon typus in continental Europe

A whale shark Rhincodon typus was found in a set-net in southern Portugal in October 2011. This was the first record for continental Europe and represents an increase of the species' known range. Upwelling events and increase in sea-surface temperatures are possible reasons for this occurrence.     

Association of the whale shark Rhincodon typus with the cannonball jellyfish Stomolophus meleagris

This study describes the first record of the whale shark Rhincodon typus association with the cannonball jellyfish Stomolophus meleagris in the Gulf of California, Mexico. Whale sharks were observed swimming and feeding among swarms of jellyfish, suggesting competition and predatory behaviour given the overlap in food preferences between both species. This finding is relevant because of the species‐wide distribution and the importance of these interactions, which should be considered in conservation strategies of R. typus and management of cannonball jellyfish fisheries.     

Northernmost record of a whale shark Rhincodon typus from the Sea of Okhotsk

The whale shark Rhincodon typus is the world's largest fish and it occurs in tropical, subtropical and warm temperate waters. Here, the northernmost record of R. typus is reported, when it was found in the Sea of Okhotsk for the first time. This occurrence can be explained by the unusually high sea surface temperature during the summer of 2012.     

Population genetic structure of Earth's largest fish, the whale shark (Rhincodon typus)

Large pelagic vertebrates pose special conservation challenges because their movements generally exceed the boundaries of any single jurisdiction. To assess the population structure of whale sharks (Rhincodon typus), we sequenced complete mitochondrial DNA control regions from individuals collected across a global distribution. We observed 51 single site polymorphisms and 8 regions with indels comprising 44 haplotypes in 70 individuals, with high haplotype (h = 0.974 +/- 0.008) and nucleotide diversity (pi = 0.011 +/- 0.006). The control region has the largest length variation yet reported for an elasmobranch (1143-1332 bp). Phylogenetic analyses reveal no geographical clustering of lineages and the most common haplotype was distributed globally. The absence of population structure across the Indian and Pacific basins indicates that oceanic expanses and land barriers in Southeast Asia are not impediments to whale shark dispersal. We did, however, find significant haplotype frequency differences (AMOVA, Phi(ST) = 0.107, P < 0.001) principally between the Atlantic and Indo-Pacific populations. In contrast to other recent surveys of globally distributed sharks, we find much less population subdivision and no evidence for cryptic evolutionary partitions. Discovery of the mating and pupping areas of whale sharks is key to further population genetic studies. The global pattern of shared haplotypes in whale sharks provides a compelling argument for development of broad international approaches for management and conservation of Earth's largest fish.     

Population structure of a whale shark Rhincodon typus aggregation in the Red Sea

The presence of whale sharks Rhincodon typus were recorded around Shib Habil, a small, coastal reef off the Red Sea coast of Saudi Arabia, from 2010 to 2015. A total of 267 suitable photographs resulting in the identification of 136 individuals, were documented from 305 encounters. Sharks were divided evenly between the sexes with no evidence of temporal or spatial segregation. All individuals were immature based on size estimates and, for males, juvenile clasper morphology. Scars were reported for 57% of R. typus with 15% showing evidence of propeller trauma. Estimates of population size and patterns of residency were calculated by modelling the lagged identification rate. Multiple models were run simultaneously and compared using the Akaike information criterion. An open population model was found to best represent the data and estimates a daily abundance between 15 and 34 R. typus during the aggregation season, with local residence times ranging from 4 to 44 days. Residence times away from Shib Habil range from 15 to 156 days with a permanent emigration–death rate between 0·07 and 0·58 individuals year^?1. These results are broadly similar to those from other aggregations of R. typus, although the observed sexual parity and integration found at this site is unique for the species and needs further study.     

Isolation and characterization of microsatellite loci in the whale shark (Rhincodon typus)

In preparation for a study on population structure of the whale shark (Rhincodon typus), nine species-specific polymorphic microsatellite DNA markers were developed. An initial screening of 50 individuals from Holbox Island, Mexico found all nine loci to be polymorphic, with two to 17 alleles observed per locus. Observed and expected heterozygosity per locus ranged from 0.200 to 0.826 and from 0.213 to 0.857, respectively. Neither statistically significant deviations from Hardy-Weinberg expectations nor statistically significant linkage disequilibrium between loci were observed. These microsatellite loci appear suitable for examining population structure, kinship assessment and other applications.     

The first record,tagging and release of a neonatal whale shark Rhincodon typus in Taiwan

On 27 October 2013, a Rhincodon typus was apparently chased by a group of Caranx ignobilis into nearshore waters near Green Island (Ludao), east of Taiwan. A fisherman brought it back to port where it was kept in a small sea pen until release. The R. typus was 78 cm total length, and was tagged and released on 29 October 2013.     

Deep-diving behaviour of a whale shark Rhincodon typus during long-distance movement in the western Indian Ocean

A whale shark Rhincodon typus satellite tagged off the coast of Mozambique showed a highly directional movement across the Mozambique Channel and around the southern tip of Madagascar, a minimum distance of 1200 km in 87 days. Dives to depths well into the mesopelagic and bathypelagic zones (1286 m maximum depth) were recorded in a bathymetrically non-constraining habitat. The water temperature range recorded during the fish's movement was 3·4–29·9° C.     

General descriptions of the dermis structure of a juvenile whale shark Rhincodon typus from the Gulf of California

The aim of this study is to provide preliminary observations on the microanatomy of Rhincodon typus skin using histology and electron microscopy analyses. Skin biopsies were obtained from a deceased juvenile male shark (548 cm total length) stranded in La Paz, Mexico, during February 2018. The results of this study evidenced the basic structure of the dermal denticles in the epidermis of the trunk of the shark, as well as the composition of the connective tissue in the hypodermis. Histological images of the hypodermis showed a high concentration of collagen fibres, formed by a large number of fine and wavy fibres of compact shape and little intercellular substance.     

Occurrences of whale shark (Rhincodon typus Smith, 1828) in the Saint Peter and Saint Paul archipelago, Brazil

The Saint Peter and Saint Paul Archipelago in the central tropical Atlantic, is an important ground of whale sharks that are commonly sighted throughout the year close to the fishing boats in the adjacencies of the islands. In sightings reported between February 2000 and November 2005, the lengths of the individuals ranged between 1.8 to 14.0 m. The causes of these concentrations in the archipelago are still unclear, once there are no upwellings and plankton concentrations for feeding, and no reproductive activities were reported. Nevertheless, they could be associated to the spawning period of the abundant flying fishes, mainly in the first semester, when sightings were more frequent.     

Seasonal trends in whale shark Rhincodon typus sightings in an established tourism site in the Gulf of California,Mexico

The number of individual whale shark Rhincodon typus sightings registered October 2015–March 2018 in Bahia de La Paz, Mexico, ranged from 73 to 129 animals per season (total 1662), with the majority of them identified as males. Density plot maps showed high correlations in sightings between sample seasons and revealed potential hotspot areas. Our study provides an essential baseline of information for the management of whale shark conservation and the associated touristic activities in the region.     

A preliminary survey of whale shark Rhincodon typus catch and trade in China: an emerging crisis

This study gives an account of spatial and temporal distribution of whale shark Rhincodon typus catch events in China on the basis of historical records and information obtained from interviews with fishing industry stakeholders. A total of 186 R. typus were recorded with key harvest areas identified as in Hainan and Zhejiang, and the peak catching seasons were May to June and September to October. Aspects of the R. typus trade are discussed, including products, markets and the process. The results suggest that R. typus is increasingly becoming a targeted resource in China as a consequence of fierce competition for large shark fins and an emerging local market for consumption of all body parts. Current obstacles and potential measures for sustainable exploitation and trade of R. typus are discussed.     

A rare occurrence of a whale shark (Rhincodon typus) in a contaminated estuary in the Southwest Atlantic Ocean

Environmental Biology of Fishes - An individual whale shark (Rhincodon typus) was recorded on January 15, 2020, during the marine megafauna drone-monitoring. The animal was filmed for approximately...     

A new record of whale shark Rhincodon typus in Brazilian waters: a report of association with Caranx crysos

In May 2011, a Rhincodon typus was sighted on the continental shelf of the central Brazilian coast, in the vicinity of a gas platform. During the video record, an interspecific following association was observed between a Caranx crysos school and the R. typus.     

Whale shark Rhincodon typus strandings in the Gulf of California,Mexico

The present study analyses whale shark Rhincodon typus stranding in the Gulf of California, Mexico, reported by the public, scientists, authorities and artisanal fishermen. A total of 14 strandings were documented during the period 2001–2018. The total length of R. typus ranged from 350–1,102 cm, with a sex ratio of 3.5:1 (males: females). This study highlights potential stranding areas, the need for better stranding protocols to assist R. typus recovery and the importance of samples from dead animals for scientific research.     

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.