Fabrice R. A. Jaine

Biology, ecology and conservation of the Mobulidae

The Mobulidae are zooplanktivorous elasmobranchs comprising two recognized species of manta rays (Manta spp.) and nine recognized species of devil rays (Mobula spp.). They are found circumglobally in tropical, subtropical and temperate coastal waters. Although mobulids have been recorded for over 400 years, critical knowledge gaps still compromise the ability to assess the status of these species. On the basis of a review of 263 publications, a comparative synthesis of the biology and ecology of mobulids was conducted to examine their evolution, taxonomy, distribution, population trends, movements and aggregation, reproduction, growth and longevity, feeding, natural mortality and direct and indirect anthropogenic threats. There has been a marked increase in the number of published studies on mobulids since c. 1990, particularly for the genus Manta, although the genus Mobula remains poorly understood. Mobulid species have many common biological characteristics although their ecologies appear to be species-specific, and sometimes region-specific. Movement studies suggest that mobulids are highly mobile and have the potential to rapidly travel large distances. Fishing pressure is the major threat to many mobulid populations, with current levels of exploitation in target fisheries unlikely to be sustainable. Advances in the fields of population genetics, acoustic and satellite tracking, and stable-isotope and fatty-acid analyses will provide new insights into the biology and ecology of these species. Future research should focus on the uncertain taxonomy of mobulid species, the degree of overlap between their large-scale movement and human activities such as fisheries and pollution, and the need for management of inter-jurisdictional fisheries in developing nations to ensure their long-term sustainability. Closer collaboration among researchers worldwide is necessary to ensure standardized sampling and modelling methodologies to underpin global population estimates and status.

Distribution, site affinity and regional movements of the manta ray, Manta alfredi (Krefft, 1868), along the east coast of Australia

Despite the increasing tourism interest worldwide for the manta ray, Manta alfredi, very little is known about its biology and ecology. Knowledge of its distribution and movement patterns is important for conservation purposes. Here we describe the distribution, site visitation and movements of M. alfredi along the east coast of Australia. Photographic identification techniques were used to identify individual manta rays at three study sites: Lady Elliot Island, North Stradbroke Island and Byron Bay. Of 388 M. alfredi individuals identified at Lady Elliot Island, 187 (48%) were subsequently re-identified at least once at the same site. In total, 31 individuals were identified at both Lady Elliot Island and North Stradbroke Island (,380km to the south) and 4 at both Lady Elliot Island and Byron Bay (,500km to the south). Manta alfredi was present all year around at Lady Elliot Island, although in higher numbers in winter, and was mainly observed at North Stradbroke Island and Byron Bay from mid-spring to mid-autumn. This is the first report of seasonal movements and site affinity forM. alfredi ineastern Australian waters and emphasises the value of photographic identification for monitoring the occurrence of individuals.

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.

Prey density threshold and tidal influence on reef manta ray foraging at an aggregation site on the Great Barrier Reef

Large tropical and sub-tropical marine animals must meet their energetic requirements in a largely oligotrophic environment. Many planktivorous elasmobranchs, whose thermal ecologies prevent foraging in nutrient-rich polar waters, aggregate seasonally at predictable locations throughout tropical oceans where they are observed feeding. Here we investigate the foraging and oceanographic environment around Lady Elliot Island, a known aggregation site for reef manta rays Manta alfredi in the southern Great Barrier Reef. The foraging behaviour of reef manta rays was analysed in relation to zooplankton populations and local oceanography, and compared to long-term sighting records of reef manta rays from the dive operator on the island. Reef manta rays fed at Lady Elliot Island when zooplankton biomass and abundance were significantly higher than other times. The critical prey density threshold that triggered feeding was 11.2 mg m-3 while zooplankton size had no significant effect on feeding. The community composition and size structure of the zooplankton was similar when reef manta rays were feeding or not, with only the density of zooplankton changing. Higher zooplankton biomass was observed prior to low tide, and long-term (~5 years) sighting data confirmed that more reef manta rays are also observed feeding during this tidal phase than other times. This is the first study to examine prey availability at an aggregation site for reef manta rays and it indicates that they feed in locations and at times of higher zooplankton biomass.

Unique Sequence of Events Triggers Manta Ray Feeding Frenzy in the Southern Great Barrier Reef, Australia

Manta rays are classified as Vulnerable to Extinction on the IUCN Red List for Threatened Species. In Australia, a key aggregation site for reef manta rays is Lady Elliot Island (LEI) on the Great Barrier Reef, ~7 km from the shelf edge. Here, we investigate the environmental processes that triggered the largest manta ray feeding aggregation yet observed in Australia, in early 2013. We use MODIS sea surface temperature (SST), chlorophyll-a concentration and photic depth data, together with in situ data, to show that anomalous river discharges led to high chlorophyll (anomalies: 10–15 mg∙m−3) and turbid (photic depth anomalies: −15 m) river plumes extending out to LEI, and that these became entrained offshore around the periphery of an active cyclonic eddy. Eddy dynamics led to cold bottom intrusions along the shelf edge (6 °C temperature decrease), and at LEI (5 °C temperature decrease). Strongest SST gradients (>1 °C∙km−1) were at the convergent frontal zone between the shelf and eddy-influenced waters, directly overlying LEI. Here, the front intensified on the spring ebb tide to attract and shape the aggregation pattern of foraging manta rays. Future research could focus on mapping the probability and persistence of these ecologically significant frontal zones via remote sensing to aid the management and conservation of marine species.

Australia’s continental-scale acoustic tracking database and its automated quality control process

Our ability to predict species responses to environmental changes relies on accurate records of animal movement patterns. Continental-scale acoustic telemetry networks are increasingly being established worldwide, producing large volumes of information-rich geospatial data. During the last decade, the Integrated Marine Observing System’s Animal Tracking Facility (IMOS ATF) established a permanent array of acoustic receivers around Australia. Simultaneously, IMOS developed a centralised national database to foster collaborative research across the user community and quantify individual behaviour across a broad range of taxa. Here we present the database and quality control procedures developed to collate 49.6 million valid detections from 1891 receiving stations. This dataset consists of detections for 3,777 tags deployed on 117 marine species, with distances travelled ranging from a few to thousands of kilometres. Connectivity between regions was only made possible by the joint contribution of IMOS infrastructure and researcher-funded receivers. This dataset constitutes a valuable resource facilitating meta-analysis of animal movement, distributions, and habitat use, and is important for relating species distribution shifts with environmental covariates.

A historical and contemporary consideration of the diet of the reef manta ray (Manta alfredi) from the Great Barrier Reef, Australia

The preserved stomach contents from the manta ray, collected in 1935, that provided the basis for the 2009 taxonomic resurrection of the species Manta alfredi, were examined. The majority of the material comprised calanoid copepods (61.7%) and trypanorhynch cestodes (34.6%), with minor contributions by arrow worms, a barnacle larva and a nematode. Comparison of the size-frequency distributions of stomach contents with that of zooplankton from the Great Barrier Reef region suggest that this manta ray preferentially ingested large copepods, or that the filter mechanism used to extract prey from the water was selective for prey items over 0.8mm in length. This is the first description of the diet of M. alfredi from stomach contents, and is consistent with previous inferences about what this species consumes.

Continental-scale animal tracking reveals functional movement classes across marine taxa

Acoustic telemetry is a principle tool for observing aquatic animals, but coverage over large spatial scales remains a challenge. To resolve this, Australia has implemented the Integrated Marine Observing System’s Animal Tracking Facility which comprises a continental-scale hydrophone array and coordinated data repository. This national acoustic network connects localized projects, enabling simultaneous monitoring of multiple species over scales ranging from 100 s of meters to 1000 s of kilometers. There is a need to evaluate the utility of this national network in monitoring animal movement ecology, and to identify the spatial scales that the network effectively operates over. Cluster analyses assessed movements and residency of 2181 individuals from 92 species, and identified four functional movement classes apparent only through aggregating data across the entire national network. These functional movement classes described movement metrics of individuals rather than species, and highlighted the plasticity of movement patterns across and within populations and species. Network analyses assessed the utility and redundancy of each component of the national network, revealing multiple spatial scales of connectivity influenced by the geographic positioning of acoustic receivers. We demonstrate the significance of this nationally coordinated network of receivers to better reveal intra-specific differences in movement profiles and discuss implications for effective management.

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.

First photographic records of the giant manta ray Manta birostris off eastern Australia.

We present the first photographic evidence of the presence of the giant manta ray Manta birostris in east Australian waters. Two individuals were photographed off Montague Island in New South Wales and off the north east coast of Tasmania, during summer 2012 and 2014, respectively. These sightings confirm previous unverified reports on the species occurrence and extend the known distribution range of M. birostris to 40°S. We discuss these findings in the context of the species’ migratory behaviour, the regional oceanography along the south east Australian coastline and local productivity events.