Depth variation in benthic community response to repeated marine heatwaves on remote Central Indian Ocean reefs.

Coral reefs are increasingly impacted by climate-induced warming events. However, there is limited empirical evidence on the variation in the response of shallow coral reef communities to thermal stress across depths. Here, we assess depth-dependent changes in coral reef benthic communities following successive marine heatwaves from 2015 to 2017 across a 5-25 m depth gradient in the remote Chagos Archipelago, Central Indian Ocean. Our analyses show an overall decline in hard and soft coral cover and an increase in crustose coralline algae, sponge and reef pavement following successive marine heatwaves on the remote reef system. Our findings indicate that the changes in benthic communities in response to elevated seawater temperatures varied across depths. We found greater changes in benthic group cover at shallow depths (5-15 m) compared with deeper zones (15-25 m). The loss of hard coral cover was better predicted by initial thermal stress, while the loss of soft coral was associated with repeated thermal stress following successive warming events. Our study shows that benthic communities extending to 25 m depth were impacted by successive marine heatwaves, supporting concerns about the resilience of shallow coral reef communities to increasingly severe climate-driven warming events.

Linking variation in planktonic primary production to coral reef fish growth and condition.

Within low-nutrient tropical oceans, islands and atolls with higher primary production support higher fish biomass and reef organism abundance. External energy subsidies can be delivered onto reefs via a range of physical mechanisms. However, the influence of spatial variation in primary production on reef fish growth and condition is largely unknown. It is not yet clear how energy subsidies interact with reef depth and slope. Here we test the hypothesis that with increased proximity to deep-water oceanic nutrient sources, or at sites with shallower reef slopes, parameters of fish growth and condition will be higher. Contrary to expectations, we found no association between fish growth rate and sites with higher mean chlorophyll-a values. There were no differences in fish δ 15N or δ 13C values between depths. The relationship between fish condition and primary production was influenced by depth, driven by increased fish condition at shallow depths within a primary production 'hotspot' site. Carbon δ 13C was depleted with increasing primary production, and interacted with reef slope. Our results indicate that variable primary production did not influence growth rates in planktivorous Chromis fieldi within 10-17.5 m depth, but show site-specific variation in reef physical characteristics influencing fish carbon isotopic composition.

Atoll-dependent variation in depth zonation of benthic communities on remote reefs.

The distribution and organisation of benthic organisms on tropical reefs are typically heterogenous yet display distinct zonation patterns across depth gradients. However, there are few datasets which inform our understanding of how depth zonation in benthic community composition varies spatially among and within different reef systems. Here, we assess the depth zonation in benthic forereef slope communities in the Central Indian Ocean, prior to the back-to-back bleaching events in 2014-2017. We compare benthic communities between shallow (5-10 m) and deep (20-25 m) sites, at two spatial scales: among and within 4 atolls. Our analyses showed the variation in both major functional groups and hard coral assemblages between depth varied among atolls, and within-atoll comparisons revealed distinct differences between shallow and deep forereef slope communities. Indicator taxa analyses characterising the hard coral community between depths revealed a higher number of coral genera characteristic of the deep forereef slopes (10) than the shallow forereef slopes (6). Only two coral genera consistently associated with both depths across all atolls, and these were Acropora and Porites. Our results reveal spatial variation in depth zonation of benthic communities, potentially driven by biophysical processes varying across depths and atolls, and provide a baseline to understand and measure the impacts of future global climate change on benthic communities across depths.

Limited impact of an invasive oyster on intertidal assemblage structure and biodiversity: the importance of environmental context and functional equivalency with native species.

Impacts of invasive species are context dependent and linked to the ecosystem they occur within. To broaden the understanding of the impact of a globally widespread invasive oyster, Crassostrea (Magallana) gigas, intertidal surveys were carried out at 15 different sites in Europe. The impact of C. gigas on macro- (taxa surrounding oyster > 1 cm) and epifaunal (taxa on oyster < 1 cm) benthic communities and α and ß-diversity was assessed and compared to those associated with native ecosystem engineers, including the flat oyster Ostrea edulis. Whilst the effect of C. gigas on benthic community structures was dependent on habitat type, epifaunal communities associated with low densities of O. edulis and C. gigas did not differ and changes in benthic assemblage structure owing to the abundance of C. gigas were therefore attributed to the presence of oyster shells. Macrofaunal α-diversity increased with C. gigas cover in muddy habitats, while epifaunal α-diversity decreased at greater oyster densities. Macrofaunal ß-diversity was greatest at low densities of C. gigas; however, it did not differ between samples without and increased densities of oysters. In contrast, epifaunal ß-diversity decreased with increasing oyster cover. Different environmental contexts enabled more independent predictions of the effect of C. gigas on native communities. These were found to be low and more importantly not differing from O. edulis. This indicates that, at low densities, C. gigas may be functionally equivalent to the declining native oyster in terms of biodiversity facilitation and aid in re-establishing benthic communities on shores where O. edulis has become extinct.

Recreational Diving Impacts on Coral Reefs and the Adoption of Environmentally Responsible Practices within the SCUBA Diving Industry.

Recreational diving on coral reefs is an activity that has experienced rapidly growing levels of popularity and participation. Despite providing economic activity for many developing coastal communities, the potential role of dive impacts in contributing to coral reef damage is a concern at heavily dived locations. Management measures to address this issue increasingly include the introduction of programmes designed to encourage environmentally responsible practices within the dive industry. We examined diver behaviour at several important coral reef dive locations within the Philippines and assessed how diver characteristics and dive operator compliance with an environmentally responsible diving programme, known as the Green Fins approach, affected reef contacts. The role of dive supervision was assessed by recording dive guide interventions underwater, and how this was affected by dive group size. Of the 100 recreational divers followed, 88 % made contact with the reef at least once per dive, with a mean (±SE) contact rate of 0.12 ± 0.01 per min. We found evidence that the ability of dive guides to intervene and correct diver behaviour in the event of a reef contact decreases with larger diver group sizes. Divers from operators with high levels of compliance with the Green Fins programme exhibited significantly lower reef contact rates than those from dive operators with low levels of compliance. The successful implementation of environmentally responsible diving programmes, which focus on influencing dive industry operations, can contribute to the management of human impacts on coral reefs.

Diet reconstruction and resource partitioning of a Caribbean marine mesopredator using stable isotope bayesian modelling.

The trophic ecology of epibenthic mesopredators is not well understood in terms of prey partitioning with sympatric elasmobranchs or their effects on prey communities, yet the importance of omnivores in community trophic dynamics is being increasingly realised. This study used stable isotope analysis of (15)N and (13)C to model diet composition of wild southern stingrays Dasyatis americana and compare trophic niche space to nurse sharks Ginglymostoma cirratum and Caribbean reef sharks Carcharhinus perezi on Glovers Reef Atoll, Belize. Bayesian stable isotope mixing models were used to investigate prey choice as well as viable Diet-Tissue Discrimination Factors for use with stingrays. Stingray δ(15)N values showed the greatest variation and a positive relationship with size, with an isotopic niche width approximately twice that of sympatric species. Shark species exhibited comparatively restricted δ(15)N values and greater δ(13)C variation, with very little overlap of stingray niche space. Mixing models suggest bivalves and annelids are proportionally more important prey in the stingray diet than crustaceans and teleosts at Glovers Reef, in contrast to all but one published diet study using stomach contents from other locations. Incorporating gut contents information from the literature, we suggest diet-tissue discrimination factors values of Δ(15)N ≈ 2.7‰ and Δ(13)C ≈ 0.9‰ for stingrays in the absence of validation experiments. The wide trophic niche and lower trophic level exhibited by stingrays compared to sympatric sharks supports their putative role as important base stabilisers in benthic systems, with the potential to absorb trophic perturbations through numerous opportunistic prey interactions.

Acoustic detection probability of bottlenose dolphins, Tursiops truncatus, with static acoustic dataloggers in Cardigan Bay, Wales.

Acoustic dataloggers are used for monitoring the occurrence of cetaceans and can aid in fulfilling statutory monitoring requirements of protected species. Although useful for long-term monitoring, their spatial coverage is restricted, and for many devices the effective detection distance is not specified. A generalized additive mixed model (GAMM) was used to investigate the effects of (1) distance from datalogger, (2) animal behavior (feeding and traveling), and (3) group size on the detection probability of bottlenose dolphins (Tursiops truncatus) with autonomous dataloggers (C-PODs) validated with visual observations. The average probability of acoustic detection for minutes with a sighting was 0.59 and the maximum detection distance ranged from 1343-1779 m. Minutes with feeding activity had higher acoustic detection rates and longer average effective detection radius (EDR) than traveling ones. The detection probability for single dolphins was significantly higher than for groups, indicating that their acoustic behavior may differ from those of larger groups in the area, making them more detectable. The C-POD is effective at detecting dolphin presence but the effects of behavior and group size on detectability create challenges for estimating density from detections as higher detection rate of feeding dolphins could yield erroneously high density estimates in feeding areas.

Thresher sharks use tail-slaps as a hunting strategy.

The hunting strategies of pelagic thresher sharks (Alopias pelagicus) were investigated at Pescador Island in the Philippines. It has long been suspected that thresher sharks hunt with their scythe-like tails but the kinematics associated with the behaviour in the wild are poorly understood. From 61 observations recorded by handheld underwater video camera between June and October 2010, 25 thresher shark shunting events were analysed. Thresher sharks employed tail-slaps to debilitate sardines at all times of day. Hunting events comprised preparation, strike, wind-down recovery and prey item collection phases, which occurred sequentially. Preparation phases were significantly longer than the others, presumably to enable a shark to windup a tail-slap. Tail-slaps were initiated by an adduction of the pectoral fins, a manoeuvre that changed a thresher shark's pitch promoting its posterior region to lift rapidly, and stall its approach. Tail-slaps occurred with such force that they may have caused dissolved gas to diffuse out of the water column forming bubbles. Thresher sharks were able to consume more than one sardine at a time, suggesting that tail-slapping is an effective foraging strategy for hunting schooling prey. Pelagic thresher sharks appear to pursue sardines opportunistically by day and night, which may make them vulnerable to fisheries. Alopiids possess specialist pectoral and caudal fins that are likely to have evolved, at least in part, for tail-slapping. The evidence is now clear; thresher sharks really do hunt with their tails.

Oceanic sharks clean at coastal seamount.

Interactions between pelagic thresher sharks (Alopias pelagicus) and cleaner wrasse were investigated at a seamount in the Philippines. Cleaning associations between sharks and teleosts are poorly understood, but the observable interactions seen at this site may explain why these mainly oceanic sharks regularly venture into shallow coastal waters where they are vulnerable to disturbance from human activity. From 1,230 hours of observations recorded by remote video camera between July 2005 and December 2009, 97 cleaner-thresher shark events were analyzed, 19 of which were interrupted. Observations of pelagic thresher sharks interacting with cleaners at the seamount were recorded at all times of day but their frequency declined gradually from morning until evening. Cleaners showed preferences for foraging on specific areas of a thresher shark's body. For all events combined, cleaners were observed to conduct 2,757 inspections, of which 33.9% took place on the shark's pelvis, 23.3% on the pectoral fins, 22.3% on the caudal fin, 8.6% on the body, 8.3% on the head, 2.1% on the dorsal fin, and 1.5% on the gills respectively. Cleaners did not preferentially inspect thresher sharks by time of day or by shark sex, but there was a direct correlation between the amount of time a thresher shark spent at a cleaning station and the number of inspections it received. Thresher shark clients modified their behavior by "circular-stance-swimming," presumably to facilitate cleaner inspections. The cleaner-thresher shark association reflected some of the known behavioral trends in the cleaner-reef teleost system since cleaners appeared to forage selectively on shark clients. Evidence is mounting that in addition to acting as social refuges and foraging grounds for large visiting marine predators, seamounts may also support pelagic ecology by functioning as cleaning stations for oceanic sharks and rays.

Early development and acquisition of Zooxanthellae in the temperate symbiotic sea anemone Anthopleura ballii (Cocks).

The ova of Anthopleura ballii become infected with zooxanthellae (endosymbiotic dinoflagellates) of maternal origin just prior to spawning. After fertilization, the zygotes undergo radial, holoblastic cleavage, and then gastrulate by invagination to form ciliated planulae. Because the zooxanthellae are localized on one side of the ovum-and later, within the blastomeres at one end of the embryo-invagination leads to the zooxanthellae being restricted to the planular endoderm and hence to the gastrodermal cells of the adult anemone. We propose that maternal inheritance of zooxanthellae plays an important part in the success of these temperate sea anemones, which live in regions where potential sources of zooxanthellae are scarce.