Alex D. Rogers

Winners and losers in a world where the high seas is closed to fishing

Fishing takes place in the high seas and Exclusive Economic Zones (EEZs) of maritime countries. Closing the former to fishing has recently been proposed in the literature and is currently an issue of debate in various international fora. We determine the degree of overlap between fish caught in these two areas of the ocean, examine how global catch might change if catches of straddling species or taxon groups increase within EEZs as a result of protection of adjacent high seas; and identify countries that are likely to gain or lose in total catch quantity and value following high-seas closure. We find that <0.01% of the quantity and value of commercial fish taxa are obtained from catch taken exclusively in the high seas, and if the catch of straddling taxa increases by 18% on average following closure because of spillover, there would be no loss in global catch. The Gini coefficient, which measures income inequality, would decrease from 0.66 to 0.33. Thus, closing the high seas could be catch-neutral while inequality in the distribution of fisheries benefits among the worlds maritime countries could be reduced by 50%.

A reversal of fortunes: climate change ‘winners’ and ‘losers’ in Antarctic Peninsula penguins

Climate change is a major threat to global biodiversity. Antarctic ecosystems are no exception. Investigating past species responses to climatic events can distinguish natural from anthropogenic impacts. Climate change produces ‘winners’, species that benefit from these events and ‘losers’, species that decline or become extinct. Using molecular techniques, we assess the demographic history and population structure of Pygoscelis penguins in the Scotia Arc related to climate warming after the Last Glacial Maximum (LGM). All three pygoscelid penguins responded positively to post-LGM warming by expanding from glacial refugia, with those breeding at higher latitudes expanding most. Northern (Pygoscelis papua papua) and Southern (Pygoscelis papua ellsworthii) gentoo sub-species likely diverged during the LGM. Comparing historical responses with the literature on current trends, we see Southern gentoo penguins are responding to current warming as they did during post-LGM warming, expanding their range southwards. Conversely, Adélie and chinstrap penguins are experiencing a ‘reversal of fortunes’ as they are now declining in the Antarctic Peninsula, the opposite of their response to post-LGM warming. This suggests current climate warming has decoupled historic population responses in the Antarctic Peninsula, favoring generalist gentoo penguins as climate change ‘winners’, while Adélie and chinstrap penguins have become climate change ‘losers’.

Too much of a good thing: sea ice extent may have forced emperor penguins into refugia during the last glacial maximum

The relationship between population structure and demographic history is critical to understanding microevolution and for predicting the resilience of species to environmental change. Using mitochondrial DNA from extant colonies and radiocarbon-dated subfossils, we present the first microevolutionary analysis of emperor penguins (Aptenodytes forsteri) and show their population trends throughout the last glacial maximum (LGM, 19.5-16 kya) and during the subsequent period of warming and sea ice retreat. We found evidence for three mitochondrial clades within emperor penguins, suggesting that they were isolated within three glacial refugia during the LGM. One of these clades has remained largely isolated within the Ross Sea, while the two other clades have intermixed around the coast of Antarctica from Adélie Land to the Weddell Sea. The differentiation of the Ross Sea population has been preserved despite rapid population growth and opportunities for migration. Low effective population sizes during the LGM, followed by a rapid expansion around the beginning of the Holocene, suggest that an optimum set of sea ice conditions exist for emperor penguins, corresponding to available foraging area.

High prevalence of obligate coral-dwelling decapods on dead corals in the Chagos Archipelago, central Indian Ocean

Small and cryptic organisms that live within the interstices of reef habitats contribute greatly to coral reef biodiversity, but are poorly studied. Many species of cryptofauna have seemingly obligate associations with live coral and are therefore considered to be very vulnerable to coral mortality. Here we report the unanticipated prevalence of obligate coral-dwelling decapod crustaceans on dead colonies of branching corals in the Chagos Archipelago (British Indian Ocean Territory) in the central Indian Ocean. A total of 205 obligate coral-dwelling decapods, including Trapezia crabs, were recorded from 43 (out of 54) dead coral colonies of Acropora and Pocillopora collected across five different atolls. Trapezia individuals found on dead corals were mainly juveniles, and the few adults were almost exclusively male. Among the shrimps (Pontoniinae), however, it was predominantly adult females found on dead corals. Obligate coral-dwelling species that typically occur only on live Pocillopora hosts (e.g., Trapezia spp.) were recorded on dead Acropora. These findings suggests that these obligate coral-dwelling decapods are not simply persisting on coral hosts that have died, but may be explicitly recruiting to or moving to dead coral hosts at certain stages in their life cycle. Variation in the abundance of live coral among sites had no affect on the presence or abundance of obligate coral-dwelling decapods on dead corals. This study shows that habitat associations of obligate coral-dwelling organisms, and their reliance on different habitat types, are complex and further work is required to establish their vulnerability to widespread habitat degradation on coral reefs.

Evolutionary dynamics of a common sub-Antarctic octocoral family.

Sequence data were obtained for five different loci, both mitochondrial (cox1, mtMutS, 16S) and nuclear (18S, 28S rDNA), from 64 species representing 25 genera of the common deep-sea octocoral family Primnoidae. We tested the hypothesis that Primnoidae have an Antarctic origin, as this is where they currently have high species richness, using Maximum likelihood and Bayesian inference methods of phylogenetic analysis. Using a time-calibrated molecular phylogeny we also investigated the time of species radiation in sub-Antarctic Primnoidae. Our relatively wide taxon sampling and phylogenetic analysis supported Primnoidae as a monophyletic family. The base of the well-supported phylogeny was Pacific in origin, indicating Primnoidae sub-Antarctic diversity is a secondary species radiation. There is also evidence for a subsequent range extension of sub-Antarctic lineages into deep-water areas of the Indian and Pacific Oceans. Conservative and speculative fossil-calibration analyses resulted in two differing estimations of sub-Antarctic species divergence times. Conservative analysis suggested a sub-Antarctic species radiation occurred ∼52MYA (95% HPD: 36-73MYA), potentially before the opening of the Drake Passage and Antarctic Circumpolar Current (ACC) formation (41-37MYA). Speculative analysis pushed this radiation back into the late Jurassic, 157MYA (95% HPD: 118-204MYA). Genus-level groupings were broadly supported in this analysis with some notable polyphyletic exceptions: Callogorgia, Fanellia, Primnoella, Plumarella, Thouarella. Molecular and morphological evidence supports the placement of Tauroprimnoa austasensis within Dasystenella and Fannyella kuekenthali within Metafannyella.

Introduction to the special issue: The global state of the ocean; interactions between stresses, impacts and some potential solutions. Synthesis papers from the International Programme on the State of the Ocean 2011 and 2012 workshops.

This Special Issue publishes papers on ocean stresses, impacts and solutions that underpinned the findings of workshops hosted by The International Programme on the State of the Ocean (IPSO; http://www.stateoftheocean.org) in partnership with the International Union for the Conservation of Nature (IUCN: http://http:// www.iucn.org/). IPSO was founded to investigate anthropogenic stressors and impacts on the global ocean and to define workable solutions to reduce or eliminate these problems. The distinguishing feature of this programme is that it treats the effects of such stressors collectively, taking a holistic view of marine ecosystems and impacts on them. Recent research has emphasised that to assess the totality of human impacts on the oceans, and the biodiversity it contains, and the resultant negative effects on the goods and services provided by marine ecosystems, the interactions between stresses must be resolved. This is critical because many direct and indirect human stressors act in a cumulative or synergistic fashion. A well-known example of this is the over exploitation of algivorous fish species on coral reefs leading to a decrease in reef resilience, with respect to shocks such as mass coral bleaching, and promoting the phase shift from coral-dominated to algal dominated systems (e.g. Hughes, 1994; Mumby et al., 2006; Hoegh-Guldberg et al., 2007; Hughes et al., 2007). Another example is the increasing recognition of interactions between overfishing and nutrient pollution (eutrophication) in causing cascading changes in marine ecosystems via food-web interactions (e.g. Daskalov, 2002; Daskalov et al., 2007). Climate-change effects, including ocean warming, acidification and hypoxia all potentially interact with each other and with other human impacts including overfishing, pollution and the establishment of invasive species (e.g. Cheung et al., 2010; Johnson et al., 2011; Doney et al., 2012). Addressing the direct and indirect human impacts on the ocean requires a holistic approach to develop viable and practical approaches to reduce or eliminate current degradation of marine ecosystems. Such approaches must be joined up, for example, the effectiveness of local action to reduce direct human stresses on coral reefs must come with global-level actions to reduce CO2 emissions. Efforts to reduce fishing mortality to a point where fishing should become sustainable (i.e. below MSY) is of little use if essential fish habitat is eliminated by destructive fishing methods and the resilience of the ecosystem eroded through bycatch of nontarget species. Such an approach, which aims to maintain ecosystem health whilst enabling the provision of the goods and services humankind requires, forms the basis for ecosystem-based management (Thrush and Dayton, 2010). It is important to also bear in

Low connectivity between 'scaly-foot gastropod' (Mollusca: Peltospiridae) populations at hydrothermal vents on the Southwest Indian Ridge and the Central Indian Ridge

Hydrothermal vents on mid-oceanic ridges are patchily distributed and host many taxa endemic to deep-sea chemosynthetic environments, whose dispersal may be constrained by geographical barriers. The aim of this study was to investigate the connectivity of three populations of the ‘scaly-foot gastropod’ (Chrysomallon squamiferum Chen et al., 2015), a species endemic to hydrothermal vents in the Indian Ocean, amongst two vent fields on the Central Indian Ridge (CIR) and Longqi field, the first sampled vent field on the Southwest Indian Ridge (SWIR). Connectivity and population structure across the two mid-oceanic ridges were investigated using a 489-bp fragment of the cytochrome oxidase c subunit I (COI) gene. Phylogeographical approaches used include measures of genetic differentiation (FST), reconstruction of parsimony haplotype network, mismatch analyses and neutrality tests. Relative migrants per generation were estimated between the fields. Significant differentiation (FST = 0.28–0.29, P < 0.001) was revealed between the vent field in SWIR and the two in CIR. Signatures were detected indicating recent bottleneck events followed by demographic expansion in all populations. Estimates of relative number of migrants were relatively low between the SWIR and CIR, compared with values between the CIR vent fields. The present study is the first to investigate connectivity between hydrothermal vents across two mid-ocean ridges in the Indian Ocean. The phylogeography revealed for C. squamiferum indicates low connectivity between SWIR and CIR vent populations, with implications for the future management of environmental impacts for seafloor mining at hydrothermal vents in the region, as proposed for Longqi.

Phylogeny and systematics of deep-sea sea pens (Anthozoa: Octocorallia: Pennatulacea)

Molecular methods have been used for the first time to determine the phylogeny of families, genera and species within the Pennatulacea (sea pens). Variation in ND2 and mtMutS mitochondrial protein-coding genes proved adequate to resolve phylogenetic relationships among pennatulacean families. The gene mtMutS is more variable than ND2 and differentiates all genera, and many pennatulacean species. A molecular phylogeny based on a Bayesian analysis reveals that suborder Sessiliflorae is paraphyletic and Subselliflorae is polyphyletic. Many families of pennatulaceans do not represent monophyletic groups including Umbellulidae, Pteroeididae, and Kophobelemnidae. The high frequency of morphological homoplasy in pennatulaceans has led to many misinterpretations in the systematics of the group. The traditional classification scheme for pennatulaceans requires revision.

Body Size Versus Depth

Body size (weight per individual) is an important concept in ecology. It has been studied in the deep sea where a decrease in size with increasing depth has often been found. This has been explained as an adaptation to food limitation where size reduction results in a lowered metabolic rate and a decreased energetic requirement. However, observations vary, with some studies showing an increase in size with depth, and some finding no depth correlation at all. Here, we collected data from peer-reviewed studies on macro- and meiofaunal abundance and biomass, creating two datasets allowing statistical comparison of factors expected to influence body size in meio- and macrofaunal organisms. Our analyses examined the influence of region, taxonomic group and sampling method on the body size of meiofauna and macrofauna in the deep sea with increasing depth, and the resulting models are presented. At the global scale, meio- and macrofaunal communities show a decrease in body size with increasing depth as expected with the food limitation hypothesis. However, at the regional scale there were differences in trends of body size with depth, either showing a decrease (e.g. southwest Pacific Ocean; meio- and macrofauna) or increase (e.g. Gulf of Mexico; meiofauna only) compared to a global mean. Taxonomic groups also showed differences in body size trends compared to total community average (e.g. Crustacea and Bivalvia). Care must be taken when conducting these studies, as our analyses indicated that sampling method exerts a significant influence on research results. It is possible that differences in physiology, lifestyle and life history characteristics result in different responses to an increase in depth and/or decrease in food availability. This will have implications in the future as food supply to the deep sea changes as a result of climate change (e.g. increased ocean stratification at low to mid latitudes and reduced sea ice duration at high latitudes).

Body Size Versus Depth: Regional and Taxonomical Variation in Deep-Sea Meio- and Macrofaunal Organisms

Body size (weight per individual) is an important concept in ecology. It has been studied in the deep sea where a decrease in size with increasing depth has often been found. This has been explained as an adaptation to food limitation where size reduction results in a lowered metabolic rate and a decreased energetic requirement. However, observations vary, with some studies showing an increase in size with depth, and some finding no depth correlation at all. Here, we collected data from peer-reviewed studies on macro- and meiofaunal abundance and biomass, creating two datasets allowing statistical comparison of factors expected to influence body size in meio- and macrofaunal organisms. Our analyses examined the influence of region, taxonomic group and sampling method on the body size of meiofauna and macrofauna in the deep sea with increasing depth, and the resulting models are presented. At the global scale, meio- and macrofaunal communities show a decrease in body size with increasing depth as expected with the food limitation hypothesis. However, at the regional scale there were differences in trends of body size with depth, either showing a decrease (e.g. southwest Pacific Ocean; meio- and macrofauna) or increase (e.g. Gulf of Mexico; meiofauna only) compared to a global mean. Taxonomic groups also showed differences in body size trends compared to total community average (e.g. Crustacea and Bivalvia). Care must be taken when conducting these studies, as our analyses indicated that sampling method exerts a significant influence on research results. It is possible that differences in physiology, lifestyle and life history characteristics result in different responses to an increase in depth and/or decrease in food availability. This will have implications in the future as food supply to the deep sea changes as a result of climate change (e.g. increased ocean stratification at low to mid latitudes and reduced sea ice duration at high latitudes).