Christoph A. Rohner

How large is the world's largest fish? Measuring whale sharks Rhincodon typus with laser photogrammetry

Laser photogrammetry was found to be a promising new cost-effective technique for measuring free-swimming whale sharks Rhincodon typus. Photogrammetric measurements were more precise than visual size estimates by experienced researchers, with results from the two methods differing by 9· 8 ± 1· 1% (mean ±s.e.). A new metric of total length and the length between the fifth gill and first dorsal fin (r² = 0· 93) is proposed to facilitate easy, accurate length measurements of whale sharks in the field.

Stable isotope and signature fatty acid analyses suggest reef manta rays feed on demersal zooplankton.

Assessing the trophic role and interaction of an animal is key to understanding its general ecology and dynamics. Conventional techniques used to elucidate diet, such as stomach content analysis, are not suitable for large threatened marine species. Non-lethal sampling combined with biochemical methods provides a practical alternative for investigating the feeding ecology of these species. Stable isotope and signature fatty acid analyses of muscle tissue were used for the first time to examine assimilated diet of the reef manta ray Manta alfredi, and were compared with different zooplankton functional groups (i.e. near-surface zooplankton collected during manta ray feeding events and non-feeding periods, epipelagic zooplankton, demersal zooplankton and several different zooplankton taxa). Stable isotope δ15N values confirmed that the reef manta ray is a secondary consumer. This species had relatively high levels of docosahexaenoic acid (DHA) indicating a flagellate-based food source in the diet, which likely reflects feeding on DHA-rich near-surface and epipelagic zooplankton. However, high levels of ω6 polyunsaturated fatty acids and slightly enriched δ13C values in reef manta ray tissue suggest that they do not feed solely on pelagic zooplankton, but rather obtain part of their diet from another origin. The closest match was with demersal zooplankton, suggesting it is an important component of the reef manta ray diet. The ability to feed on demersal zooplankton is likely linked to the horizontal and vertical movement patterns of this giant planktivore. These new insights into the habitat use and feeding ecology of the reef manta ray will assist in the effective evaluation of its conservation needs.

Mobulid rays feed on euphausiids in the Bohol Sea

Mobulid rays have a conservative life history and are caught in direct fisheries and as by-catch. Their subsequent vulnerability to overexploitation has recently been recognized, but fisheries management can be ineffective if it ignores habitat and prey preferences and other trophic interactions of the target species. Here, we assessed the feeding ecology of four mobulids (Manta birostris, Mobula tarapacana, M. japanica, M. thurstoni) in the Bohol Sea, Philippines, using stomach contents analysis of fisheries specimens landed between November and May in 2013–2015. We show that the mobulids feed heavily on euphausiid krill while they are in the area for approximately six months of the year. We found almost no trophic separation among the mobulid species, with Euphausia diomedeae as the major prey item for all species, recorded in 81 of 89 total stomachs (91%). Mobula japanica and M. thurstoni almost exclusively had this krill in their stomach, while M. tarapacana had a squid and fish, and Ma. birostris had myctophid fishes and copepods in their stomachs in addition to E. diomedeae. This krill was larger than prey for other planktivorous elasmobranchs elsewhere and contributed a mean of 61 364 kcal per stomach (±105 032 kcal s.e., range = 0–631 167 kcal). Our results show that vertically migrating mesopelagic species can be an important food resource for large filter feeders living in tropical seas with oligotrophic surface waters. Given the conservative life history of mobulid rays, the identification of common foraging grounds that overlap with fishing activity could be used to inform future fishing effort.

Population Structure, Abundance and Movement of Whale Sharks in the Arabian Gulf and the Gulf of Oman.

Data on the occurrence of whale sharks, Rhincodon typus, in the Arabian Gulf and Gulf of Oman were collected by dedicated boat surveys and via a public-sightings scheme during the period from 2011 to 2014. A total of 422 individual whale sharks were photo-identified from the Arabian Gulf and the northern Gulf of Oman during that period. The majority of sharks (81%, n = 341) were encountered at the Al Shaheen area of Qatar, 90 km off the coast, with the Musandam region of Oman a secondary area of interest. At Al Shaheen, there were significantly more male sharks (n = 171) than females (n = 78; X2 = 17.52, P < 0.05). Mean estimated total length (TL) for sharks was 6.90 m ± 1.24 (median = 7 m; n = 296). Males (7.25 m ± 1.34; median = 8 m, n = 171) were larger than females (6.44 m ±1.09; median = 7 m, n = 78; Mann-Whitney U test, p < 0.01). Of the male sharks assessed for maturity 63% were mature (n = 81), with 50% attaining maturity by 7.29 m and 100% by 9.00 m. Two female sharks of >9 m individuals were visually assessed as pregnant. Connectivity among sharks sighted in Qatari, Omani and UAE waters was confirmed by individual spot pattern matches. A total of 13 identified sharks were re-sighted at locations other than that at which they were first sighted, including movements into and out of the Arabian Gulf through the Strait of Hormuz. Maximum likelihood techniques were used to model an estimated combined population for the Arabian Gulf and Gulf of Oman of 2837 sharks ± 1243.91 S.E. (95% C.I. 1720–6295). The Al Shaheen aggregation is thus the first site described as being dominated by mature males while the free-swimming pregnant females are the first reported from the Indian Ocean.

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.

Some like it hot: Repeat migration and residency of whale sharks within an extreme natural environment

The Arabian Gulf is the warmest sea in the world and is host to a globally significant population of the whale shark Rhincodon typus. To investigate regional whale shark behaviour and movements, 59 satellite-linked tags were deployed on whale sharks in the Al Shaheen area off Qatar from 2011–14. Four different models of tag were used throughout the study, each model able to collect differing data or quantities of data. Retention varied from one to 227 days. While all tagged sharks crossed international maritime boundaries, they typically stayed within the Arabian Gulf. Only nine sharks dispersed through the narrow Strait of Hormuz into the Gulf of Oman. Most sharks stayed close to known or suspected feeding aggregation sites over summer months, but dispersed throughout the Arabian Gulf in winter. Sharks rarely ventured into shallow areas (<40 m depth). A single, presumably pregnant female shark was the sole animal to disperse a long distance, crossing five international maritime boundaries in 37 days before the tag detached at a distance of approximately 2644 km from the tagging site, close to the Yemeni-Somali border. No clear space-use differentiation was evident between years, for sharks of different sizes, or between sexes. Whale sharks spent the most time (~66%) in temperatures of 24–30°C and in shallow waters <100 m depth (~60%). Sharks spent relatively more time in cooler (X2 = 121.692; p<0.05) and deeper (X2 = 46.402; p<0.05) water at night. Sharks rarely made dives deeper than 100 m, reflecting the bathymetric constraints of the Gulf environment. Kernel density analysis demonstrated that the tagging site at Al Shaheen was the regional hotspot for these sharks, and revealed a probable secondary aggregation site for whale sharks in nearby Saudi Arabian waters. Analysis of visual re-sightings data of tagged sharks revealed that 58% of tagged individuals were re-sighted back in Al Shaheen over the course of this study, with 40% recorded back at Al Shaheen in the year following their initial identification. Two sharks were confirmed to return to Al Shaheen in each of the five years of study.

Opecoelidae (Digenea) in northern Great Barrier Reef goatfishes (Perciformes: Mullidae)

The Opecoelidae (Digenea) of eight species of goatfishes (Perciformes: Mullidae) were studied off Lizard Island on the northern Great Barrier Reef (GBR). Host species included Mulloidichthys flavolineatus (Lacépède), M. vanicolensis (Valenciennes), Parupeneus barberinus (Lacépède), P. ciliatus (Lacépède), P. cyclostomus (Lacépède), P. indicus (Shaw), P. multifasciatus (Quoy & Gaimard), P. spilurus (Bleeker) and P. trifasciatus (Lacépède). Seven opecoelid species were found, of which Paropecoelus corneliae n. sp., Pa. leonae n. sp. and Pseudopecoeloides engeleri n. sp. are described as new. Paropecoelus elongatus (Ozaki, 1928), Pa. sogandaresi Pritchard, 1966 and Podocotyloides parupenei (Manter, 1963) are recorded and a description is provided for each. A second species of Pseudopecoeloides remains unnamed for want of sufficient morphological material. The seven species were distinguished on the basis of morphology and molecular analysis of 56 ITS2 rDNA sequences. Sequences differed by nine to 68 base pairs between species combinations recognised by morphology; no intraspecific variation was detected. Each opecoelid species infected between two and five mullid species, but none of the species has been found in non-mullid hosts. Every mullid species was infected with at least one species of opecoelid and Parupeneus ciliatus had all seven species. The Opecoelidae of GBR goatfishes are consistent with the typical pattern of stenoxenicity seen for trematodes of GBR fishes.

Satellite tagging highlights the importance of productive Mozambican coastal waters to the ecology and conservation of whale sharks

The whale shark Rhincodon typus is an endangered, highly migratory species with a wide, albeit patchy, distribution through tropical oceans. Ten aerial survey flights along the southern Mozambican coast, conducted between 2004–2008, documented a relatively high density of whale sharks along a 200 km stretch of the Inhambane Province, with a pronounced hotspot adjacent to Praia do Tofo. To examine the residency and movement of whale sharks in coastal areas around Praia do Tofo, where they may be more susceptible to gill net entanglement, we tagged 15 juveniles with SPOT5 satellite tags and tracked them for 2–88 days (mean = 27 days) as they dispersed from this area. Sharks travelled between 10 and 2,737 km (mean = 738 km) at a mean horizontal speed of 28 ± 17.1 SD km day−1. While several individuals left shelf waters and travelled across international boundaries, most sharks stayed in Mozambican coastal waters over the tracking period. We tested for whale shark habitat preferences, using sea surface temperature, chlorophyll-a concentration and water depth as variables, by computing 100 random model tracks for each real shark based on their empirical movement characteristics. Whale sharks spent significantly more time in cooler, shallower water with higher chlorophyll-a concentrations than model sharks, suggesting that feeding in productive coastal waters is an important driver of their movements. To investigate what this coastal habitat choice means for their conservation in Mozambique, we mapped gill nets during two dedicated aerial surveys along the Inhambane coast and counted gill nets in 1,323 boat-based surveys near Praia do Tofo. Our results show that, while whale sharks are capable of long-distance oceanic movements, they can spend a disproportionate amount of time in specific areas, such as along the southern Mozambique coast. The increasing use of drifting gill nets in this coastal hotspot for whale sharks is likely to be a threat to regional populations of this iconic species.

Satellite tagging of rehabilitated green sea turtles Chelonia mydas from the United Arab Emirates, including the longest tracked journey for the species

We collected movement data for eight rehabilitated and satellite-tagged green sea turtles Chelonia mydas released off the United Arab Emirates between 2005 and 2013. Rehabilitation periods ranged from 96 to 1353 days (mean = 437 ± 399 days). Seven of the eight tagged turtles survived after release; one turtle was killed by what is thought to be a post-release spear gun wound. The majority of turtles (63%) used shallow-water core habitats and established home ranges between Dubai and Abu Dhabi, the same area in which they had originally washed ashore prior to rescue. Four turtles made movements across international boundaries, highlighting that regional cooperation is necessary for the management of the species. One turtle swam from Fujairah to the Andaman Sea, a total distance of 8283 km, which is the longest published track of a green turtle. This study demonstrates that sea turtles can be successfully reintroduced into the wild after sustaining serious injury and undergoing prolonged periods of intense rehabilitation.

Life History, Growth, and Reproductive Biology of Four Mobulid Species in the Bohol Sea, Philippines

In light of the global decline of mobulid populations and the necessity for sustainable fisheries management, baseline data for population dynamics were collected from a targeted fishery in the Bohol Sea, Philippines. This study focused on life-history parameters and reproductive cycles of four mobulid rays (Mobula thurstoni, M. japanica, M. tarapacana, and Manta birostris), and re-estimated their intrinsic population growth rates. Size and reproductive data were collected from 1,509 specimens (30% of catch) landed in two fishing seasons in 2015 and 2016. Size-at-birth was reviewed, and analysis of the embryos and follicles did not show any clear seasonality in the reproductive cycle, but supported an interbreeding interval. Females of all species matured at a larger size than males, and exhibited a larger size-at-pregnancy than -at-maturity. This delay in reproduction resulted in population growth rates lower than the actual rmax when based on size-at-pregnancy (rmat=rmax=0.016-0.055 year-1 and rpreg=0.008-0.044 year-1), and a population doubling time of 15.8-86.6 years. This study suggests that population growth rates previously reported were overestimated. In light of the Convention on International Trades of Endangered Species (CITES) and Convention on Migratory Species (CMS) assessments, while fisheries management should reflect the delayed maturation of these species and the slower population growth potential, at the current status of these population, the sustainability of any exploitation level seems unrealistic and strongly discouraged.