The ecology of scattering layer biota around Indian Ocean seamounts and islands

The waters of the open ocean constitute the largest living space on Earth but despite its obvious significance to the biosphere, the open ocean remains an unexplored frontier. With a regional focus on the Indian Ocean, this thesis investigates (i) the distribution of pelagic biota on basin scales, (ii) the effect of abrupt topography on pelagic biota and their predator-prey relationships, and (iii) the use of genetic techniques to elucidate population connectivity and dispersal of pelagic taxa. (i) Pelagic scattering layers (SLs) were surveyed with scientific echosounders across the southwest (SWIO) and central Indian Ocean to investigate their vertical and geographical distribution. Structurally distinct SL regimes were found across the Subantarctic Front, and may explain recently observed foraging behaviours of southern elephant seals. Regression models indicated a close relationship between sea surface temperature and mean volume backscatter, with significantly elevated backscatter in the subtropical convergence zone. The heterogeneous distribution of scattering layer biota may have implications for predator foraging and carbon cycling in the Indian Ocean. (ii) Acoustic surveys revealed diverse interactions between SLs, aggregations and topography around islands as well as shallow (<200m) and intermediate (200-800m) seamounts at spatial scales from 1 to 100 km. Epi-and mesopelagic backscatter was increased around reefs and banks of the Chagos archipelago, indicating connectivity between oceanic and neritic systems. SWIO seamounts harboured summit-associated aggregations, but the distributions of surrounding SLs did not follow a general pattern. Downstream SL depletion was observed in one location and combined with stomach content analyses, provides an insight into the mechanics of prey flux between open-ocean and seamount ecosystems. (iii) A mitochondrial marker was used to assess the population structure and demography of the hatchetfish Argyropelecus aculeatus in the SWIO. The results are suggestive of a single, well-connected population and indicate a recent population expansion around 0.14 million years ago. This highlights that even highly abundant mesopelagic populations are vulnerable to global climatic changes. Dispersal and recruitment are key ecological processes structuring seamount communities and are directly relevant for the management of exploited populations. Genetic barcoding was evaluated as a means to identify cryptic larval specimens of eels (leptocephali) and spiny lobsters (phyllosomata). Identification success was limited, but indicated the presence of 3-4 phyllosoma clades and 5-6 leptocephalus clades along the SWIR."Additional funds came from the German National Academic Foundation, the Rehbock Fund, the Fisheries Society of the British Isles, and the Marin Alliance for Science and Technology for Scotland (MASTS) pooling initiative (Grants DSSG1 and SG98)." -- AcknowledgementsData collection in the Southwest Indian Ocean was undertaken as part of the Southwest Indian Ocean Seamounts Project (www.iucn.org/marine/seamounts) supported by the EAF Nansen Project, the Food and Agriculture Organization of the United Nations, the Global Environment Facility, the International Union for the Conservation of Nature, the Natural Environment Research Council (Grant NE/F005504/1), the Leverhulme Trust (Grant F00390C) and the Total Foundation. Shiptime in the Chagos was funded by the UK Foreign and Commonwealth Office." -- Acknowledgement

The distribution of pelagic sound scattering layers across the southwest Indian Ocean

Ship of Opportunity Data were sourced from the Integrated Marine Observing System (IMOS)—an initiative of the Australian Government being conducted as part of the National Collaborative Research Infrastructure Strategy and the Super Science Initiative. Other acoustic data were collected as part of the Southwest Indian Ocean Seamounts Project (http://www.iucn.org/marine/seamounts) which was supported supported by the EAF Nansen Project, the Food and Agriculture Organization of the United Nations, the Global Environment Facility, the International Union for the Conservation of Nature, the Natural Environment Research Council (Grant NE/F005504/1), the Leverhulme Trust (Grant F00390C) and the Total Foundation. We thank the Masters, officers, crews and science parties of cruises DFN 2009-410 and JCO66/67 for their assistance during echosounder calibration and data acquisition, and two anonymous reviewers for their comments. PHBS was supported by the German National Academic Foundation, a Cusanuswerk doctoral fellowship, and a Lesley & Charles Hilton-Brown Scholarship.Shallow and deep scattering layers (SLs) were surveyed with split-beam echosounders across the southwest Indian Ocean (SWIO) to investigate their vertical and geographical distribution. Cluster analysis was employed to objectively classify vertical backscatter profiles. Correlations between backscatter and environmental covariates were modelled using generalized additive mixed models (GAMMs) with spatial error structures. Structurally distinct SL regimes were found across the Subantarctic Front. GAMMs indicated a close relationship between sea surface temperature and mean volume backscatter, with significantly elevated backscatter in the subtropical convergence zone. The heterogeneous distribution of scattering layer biota reflects the biogeographic zonation of the survey area and is likely to have implications for predator foraging and carbon cycling in the Indian Ocean.PostprintPeer reviewe

Changing measurements or changing movements? Sampling scale and movement model identifiability across generations of biologging technology

Animal movement patterns contribute to our understanding of variation in breeding success and survival of individuals, and the implications for population dynamics. Over time, sensor technology for measuring movement patterns has improved. Although older technologies may be rendered obsolete, the existing data are still valuable, especially if new and old data can be compared to test whether a behavior has changed over time. We used simulated data to assess the ability to quantify and correctly identify patterns of seabird flight lengths under observational regimes used in successive generations of wet/dry logging technology. Care must be taken when comparing data collected at differing timescales, even when using inference procedures that incorporate the observational process, as model selection and parameter estimation may be biased. In practice, comparisons may only be valid when degrading all data to match the lowest resolution in a set. Changes in tracking technology, such as the wet/dry loggers explored here, that lead to aggregation of measurements at different temporal scales make comparisons challenging. We therefore urge ecologists to use synthetic data to assess whether accurate parameter estimation is possible for models comparing disparate data sets before planning experiments and conducting analyses such as responses to environmental changes or the assessment of management actions

Enhanced pelagic biomass around coral atolls

T.B.L. was supported by the Marine Biodiversity Hub through the Australian Government’s National Environmental Research Program (NERP). P.H.B.-S. was supported by a Cusanuswerk doctoral fellowship, a Lesley & Charles Hilton-Brown Scholarship, University of St. Andrews, and a grant from the Fisheries Society of the British Isles. M.J.C. was supported by Australian Research Council grant FS110200057.Understanding the processes driving the distribution of mid-water prey such as euphausiids and lanternfish is important for effective management and conservation. In the vicinity of abrupt topographic features such as banks, seamounts and shelf-breaks, mid-water faunal biomass is often elevated, making these sites candidates for special protection. We investigated the spatial distribution of water column acoustic backscatter - a proxy for macrozoo - plankton and fish biomass - in the 9 km transition zone between the pelagos and coral atolls in the Chagos Archipelago (6° N, 72° E). The purpose was to determine the magnitude and distance over which bathymetry may enhance biomass in the mid-water, and thereby identify the scale over which static topographic features could influence the open ocean. Two distinct sound scattering layers were identified, from the surface to 180 m and from 300 to 600 m, during daytime. Both layers exhibited significant increases in backscatter near features. Close to features, the shallow layer backscatter was ca. 100 times higher and was driven partly by increasing numbers of larger individuals, evident as single target echoes. We determine the regional scale of influence of features on pelagic biomass enhancement to be ca. 1.8 km in the Chagos Archipelago, and suggest possible ecological explanations that may support it. Our approach determining the scale of influence of bathymetry should be applied during the process of marine reserve design, in order to improve protection of mid-water fauna associated with topographical features, such as seamounts and coral reefs.PostprintPeer reviewe

Seamount influences on mid-water shrimps (Decapoda) and Gnathophausiids (Lophogastridea) of the South-West Indian ridge

This study was conducted under the UNDP/IUCN project, funded by the Global Environment Facility (GEF). The authors thank the School of Biology at the University of St Andrews and the National Environmental Research Council (NERC) for funding toward Tom B Letessier's PhD.Maintenance of often-observed elevated levels of pelagic diversity and biomass on seamounts, that are of relevance to conservation and fishery management, involves complex interactions between physical and biological variables that remain poorly understood. To untangle these biophysical processes we explore factors influencing the distribution of epi- and meso-pelagic (0–1000 m) micronektonic crustaceans (>15 mm; order Lophogastridea, family Gnathophausiidea; and order Decapoda) on and off seamounts along the South West Indian Ridge (SWIR, 27° to 42°S) and on a seamount off the Madagascar Ridge (31.6°S, 42.8°E). Thirty-one species of micronektic crustaceans were caught using mid-water trawls within the study are but there was no apparent latitude-related patterns in species richness or abundance. Species richness predicted by rarefraction curves and numerical abundance was highest in the vicinity (800 m). The dominant species assemblage comprised the shrimps Systellaspis debilis (37%) and Sergia prehensilis (34%), and was restricted to seamounts on the subtropical SWIR. Our observations suggest that the ‘oasis effect’ of seamounts conventionally associated with higher trophic levels is also applicable to pelagic micronektic crustaceans at lower trophic levels. We suggest that the enhanced biomass and species richness attributed is due to ‘habitat enrichment’, whereby seamounts provide favourable habitats for both pelagic and bentho-pelagic mid-water crustaceans.PostprintPeer reviewe

Drivers of octopus abundance and density in an anchialine lake: a 30 year comparison

Anchialine systems are isolated from the sea and often support species’ populations distinct from their marine counterparts. Sweetings Pond, an anchialine lake on the island of Eleuthera in The Bahamas was identified as a site of high Caribbean reef octopus, Octopus briareus (Robson, 1929) density, relative to coastal populations. However, observed deterioration in local benthic habitat and increased anthropogenic influence over the last 30 years imply that this octopus population may have undergone density and distribution shifts in response to these changing conditions. Here, we assess the system wide octopus density to provide an updated estimate. We hypothesize that despite depressed habitat availability in the 1980s, it will now support octopus densities less than historical estimates because of increasing human impact on the system. Drivers of abundance were also modelled, testing ecological hypotheses of the relationship between octopus count and prey, habitat coverage, and human disturbance. Octopus briareus were found in 7 of 27 of surveys with a mean survey count of 0.630 ± 1.25 per 900 m2. Octopus density did not vary significantly between sites. Octopus count was predicted to increase with increasing cover of calcareous rubble and the density of a preferred prey species, and intriguingly, counts decreased as a function of natural den abundance. System wide octopus density was comparable to earlier studies from the 1980s (1982 = 717.38 per km2; 1983 = 282.59 per km2; 2019 = 643.81 per km2) with no significant difference between years. Given the ecosystem’s unique and closed ecological community and the population dynamics and distribution drivers we present, Sweetings Pond has the potential to act as a ‘natural laboratory’ to explore further questions about marine insular systems and their influence on species populations in terms of ecological and behavioural change

An experimental assessment of social tolerance and den ecology in a high-density octopus population

Long held notions of the universally asocial octopus are being challenged due to the identification of high-density and interacting octopus populations in Australia, Indonesia, Japan and the deep sea. This study experimentally assessed the social tolerance and presence of potential prey items of Caribbean reef octopus, Octopus briareus, in a tropical marine lake (25°21′40″N, 76°30′40″W) on the island of Eleuthera, The Bahamas, by deploying artificial dens in multi-den groups or ‘units’ in the months of May and June 2019. Fifteen octopus were observed occupying dens (n = 100), resulting in 13 den units being occupied (n = 40). Two examples of adjacent occupation within a single den unit were identified but with zero examples of cohabitation/den sharing. Ecological models showed den and den unit occupation was predicted to increase with depth and differ between sites. Octopus also displayed no preference for isolated or communal units but preferred isolated dens over dens adjacent to others. Additionally, 47 % of occupied dens contained bivalve or crustacean items with no epifauna on their interior surface. The lack of epifauna suggests that these items have been recently ‘cleaned’ by occupying octopus and so represent likely prey. This study presents evidence of possible antisocial den use by O. briareus, a modification of the default ‘asocial’ ignoring of conspecifics typically attributed to octopus. This is likely in response to the high population density and may imply behavioural plasticity, making this system appropriate for further scrutiny as a research location on the influence of large, insular environments on marine species

Spatial variation in spring arrival patterns of Afro‐Palaearctic bird migration across Europe

Aim: Geographical patterns of migrant species arrival have been little studied, despite their relevance to global change responses. Here, we quantify continent-wide inter-specific variation in spatio-temporal patterns of spring arrival of 30 common migrant bird species and relate these to species characteristics and environmental conditions.Location: EuropeTime period: 2010-2019Major taxa studied: Birds, 30 speciesMethods: Using citizen science data from EuroBirdPortal, we modelled arrival phenology for 30 Afro-Palearctic migrant species across Europe to extract start and duration of species arrival at a 400 km square resolution. We related inter and intra-specific variation in arrival and duration to species characteristics and temperature at the start of the growing season (green-up) .Results: Spatial variation in start of arrival times indicates it took on average 1.6 days for the leading migratory front to move northwards by 100 km (range: 0.6—2.5 days). There was a major gradient in arrival phenology, from species which arrived earlier, least synchronously, in colder temperatures and progressed slowly northwards to species which arrived later, most synchronously and in warmer temperatures, and advanced quickly through Europe. The slow progress of early arrivers suggests that temperature limits their northward advance; this group included Aerial Insectivores and species wintering north of the Sahel. For the late arrivers, which included species wintering further south, seasonal resource availability in Africa may delay their arrival into Europe.Main conclusions: We found support for the green-wave hypothesis applying widely to migratory landbirds. Species arrival phenologies are linked to ecological differences between taxa, such as diet, and wintering location. Understanding these differences informs predictions of species’ sensitivity to global change. Publishing these arrival phenologies will facilitate further research and have additional conservation benefits such as informing designation of hunting seasons. Our methods are applicable to any taxa with repeated occurrence data across large scales. Key words: phenology, European-African migrants, bird migration, spring arrival, spatial variation, intraspecific and interspecific variation, EuroBirdPortal, citizen scientists, complete lists and casual record

The benefits of protected areas for bird population trends may depend on their condition

The Post-2020 Global Biodiversity Framework calls for at least 30% of land and sea to be protected by 2030. Whilst there is growing evidence that protected areas can benefit biodiversity, to achieve the greatest possible gains from their expansion, we must understand how protected area quality impacts upon biodiversity metrics. We used UK BTO/JNCC/RSPB Breeding Bird Survey data and protected areas condition data from national Common Standards Monitoring, to test whether improving site condition (for which there are UK policy targets) would contribute to stated policy targets to increase species' abundance. After controlling for differences in climate, land cover, and elevation, we found a positive association between the proportion of favourable habitat and bird abundance trends in the UK, while in Wales, Scotland and Northern Ireland combined, the positive effect was also significantly greater than in unfavourable habitat. Conversely, we also found a negative effect of proportion of favourable habitat on bird abundance. There was no evidence that these relationships varied between conservation status or many of the traits considered, although there was some evidence that favourable condition was beneficial for habitat specialists, cold-adapted species, and varied by habitat. Our findings suggest that improving the condition of protected areas currently in unfavourable condition, will contribute to nature recovery as measured by species' abundance trends in some circumstances. This also suggests that achieving the “30 by 30” target without ensuring those protected areas are of sufficient quality, may not be sufficient to restore biodiversity