David Rowat

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.

Transatlantic migration and deep mid-ocean diving by basking shark.

Despite being the second largest fish, basking sharks (Cetorhinus maximus) have been assumed to remain in discrete populations. Their known distribution encompasses temperate continental shelf areas, yet until now there has been no evidence for migration across oceans or between hemispheres. Here we present results on the tracks and behaviour of two basking sharks tagged off the British Isles, one of which released its tag off Newfoundland, Canada. During the sharks transit of the North Atlantic, she travelled a horizontal distance of 9589 km and reached a record depth of 1264 m. This result provides the first evidence for a link between European and American populations and indicates that basking sharks make use of deep-water habitats beyond the shelf edge.

Population abundance and apparent survival of the Vulnerable whale shark Rhincodon typus in the Seychelles aggregation

Identifying individuals through time can provide information on population size, composition, survival and growth rates. Identification using photographs of distinctive physical characteristics has been used in many species to replace conventional marker tagging. We evaluated photographic records over 7 years of Vulnerable whale sharks Rhincodon typus , at an aggregation in the Seychelles, for estimation of population size and structure. We collected 11,681 photographs of which only 1,149 were suitable for comparison using semi-automated matching software (I 3 S) of individual spot patterns behind the gills. Photo-identification showed that there was considerable loss of marker tags and enabled an estimation of the rate of tag loss. The combination of photo-identification with marker tagging identified a total of 512 individual sharks over 2001–2007. Of these, there were 115 resightings in subsequent years with two sharks identified in 2001 resighted 5 years later in 2006 and another shark sighted in 2001 resighted in 2007. Estimates of abundance using conventional open mark–recapture models for 2004–2007 were 348–488 sharks (95% confidence interval), with a high level of entry into the population by itinerants. Annual apparent survival probability was 0.343–0.781 over 2004–2007, with an average annual recapture probability of 0.201. These results are the first to suggest a highly transient population of whale sharks around the Seychelles, indicating that international or at least regional-scale conservation approaches are required.

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 worlds largest fish at multiple spatial scales.

Long-term membership of whale sharks (Rhincodon typus) in coastal aggregations in Seychelles and Djibouti

In coastal waters of several locations globally, whale sharks (Rhincodon typus) form seasonal aggregations, most of which largely comprise juvenile males of 4–8 m length. Evaluation of the period that individuals stay within these size- and age-specific groupings will clarify our understanding of the transition between life-stages in this species and how this might affect their long-term conservation. Long-term photo-identification studies in Seychelles and Djibouti provided data to evaluate this. The Seychelles aggregation had 443 individuals averaging 5.8 m identified between 2001 and 2009; however, the Djibouti aggregation comprised smaller individuals of 3.7 m mean length with 297 individuals identified between 2003 and 2010. In Seychelles, 27% of individuals identified in 2001 were seen again in 2009, while in Djibouti none of the whale sharks identified in 2003 were seen in 2010, although 13% from 2004 were. This suggests that membership periods in the Djibouti aggregation are shorter than in the other juvenile aggregations, such as in Seychelles. Continued photo-identification monitoring of other Indian Ocean aggregations might in time show the next location of these young sharks’ life-cycle and thereby allow development of informed national and regional management plans.

New records of neonatal and juvenile whale sharks (Rhincodon typus) from the Indian Ocean

The finding of neonatal whale sharks from Northern Indian Ocean waters off of Pakistan and Bangladesh and the description of several very small whale sharks from around Seychelles during the last few years are reported. These findings are discussed in relation to published reports of growth rates, the areas of occurrence and segregation by sex, and the behaviour of very young whale sharks, which are key factors in their conservation management.

The development of robust morphometric indices from accurate and precise measurements of free-swimming whale sharks using laser photogrammetry

To enable the study of population dynamics of wild animals the determination of the age, growth rate and maturity status of a sample of the individuals present is required; consequently, obtaining repeated accurate and precise total length (TL) measurements for individuals over time can be especially valuable. However, there are limited easily applied methods to ascertain the TL of large free-swimming fish, especially the largest extant species of fish, the whale shark ( Rhincodon typus ). This study expands on previous work and presents the results of a robust laser photogrammetry system developed to achieve accurate TL, pre-caudal length (PCL) and further morphometric measurements of whale sharks observed between 2009 and 2011 in seasonal feeding aggregations located in the Seychelles and Djibouti. Calculations for repeatability (r) indicated a high level of precision for the system with r approaching 1 for both TL and PCL, increasing further with the use of morphometric measurements. TL measurements of ‘straight sample sharks’ also provided geometric mean linear regression equations to enable the prediction of TL from defined morphological indices. Continuous validation of the system against objects of a fixed length also indicated a high level of accuracy for the method of measurement. We concluded that the laser photogrammetry system can be confidently employed to obtain accurate in-water TL, PCL and morphometric measurements for R. typus , with wide ranging implications and applications for the study of R. typus , and other large marine fauna.

Undersea Constellations: The Global Biology of an Endangered Marine Megavertebrate Further Informed through Citizen Science

The whale shark is an ideal flagship species for citizen science projects because of its charismatic nature, its size, and the associated ecotourism ventures focusing on the species at numerous coastal aggregation sites. An online database of whale shark encounters, identifying individuals on the basis of their unique skin patterning, captured almost 30,000 whale shark encounter reports from 1992 to 2014, with more than 6000 individuals identified from 54 countries. During this time, the number of known whale shark aggregation sites (hotspots) increased from 13 to 20. Examination of photo-identification data at a global scale revealed a skewed sex-ratio bias toward males (overall, more than 66%) and high site fidelity among individuals, with limited movements of sharks between neighboring countries but no records confirming large, ocean basin-scale migrations. Citizen science has been vital in amassing large spatial and temporal data sets to elucidate key aspects of whale shark life history and demographics and will continue to provide substantial long-term value.

The ecological connectivity of whale shark aggregations in the Indian Ocean: a photo-identification approach

Genetic and modelling studies suggest that seasonal aggregations of whale sharks (Rhincodon typus) at coastal sites in the tropics may be linked by migration. Here, we used photo-identification (photo-ID) data collected by both citizen scientists and researchers to assess the connectedness of five whale shark aggregation sites across the entire Indian Ocean at timescales of up to a decade. We used the semi-automated program I3S (Individual Interactive Identification System) to compare photographs of the unique natural marking patterns of individual whale sharks collected from aggregations at Mozambique, the Seychelles, the Maldives, Christmas Island (Australia) and Ningaloo Reef (Australia). From a total of 6519 photos, we found no evidence of connectivity of whale shark aggregations at ocean-basin scales within the time frame of the study and evidence for only limited connectivity at regional (hundreds to thousands of kilometres) scales. A male whale shark photographed in January 2010 at Mozambique was resighted eight months later in the Seychelles and was the only one of 1724 individuals in the database to be photographed at more than one site. On average, 35% of individuals were resighted at the same site in more than one year. A Monte Carlo simulation study showed that the power of this photo-ID approach to document patterns of emigration and immigration was strongly dependent on both the number of individuals identified in aggregations and the size of resident populations.

iDNA at sea: recovery of whale shark (Rhincodon typus) mitochondrial DNA sequences from the whale shark copepod (Pandarus rhincodonicus) confirms global population structure

Funding for this study was provided by the SeaWorld Research and Rescue Foundation, the Save our Seas Foundation, the Monash University Malaysia Tropical Medicine and Biology Platform, Monash University Malaysia School of Science, Quadrant Energy, Australian Institute of Marine Science, The Western Australian Department of Environment and Conservation, The Four Seasons Resorts Maldives. Funding for SP came from Aqua-Firma, the Shark Foundation, the Save Our Seas Foundation and two private trusts.