Human-driven Benthic Jellyfish Blooms: Causes and Consequences for Coastal Marine Ecosystems

Coastal marine ecosystems are among the most impacted globally, attributable to individual and cumulative effects of human disturbance. Anthropogenic nutrient loading is one stressor that commonly affects nearshore ecosystems, including seagrass beds, and has positive and negative effects on the structure and function of coastal systems. An additional, previously unexplored mechanistic pathway through which nutrients may indirectly influence nearshore systems is by driving blooms of benthic jellyfish. My dissertation research, conducted on Abaco Island, Bahamas, focused on elucidating the role that benthic jellyfish have in structuring systems in which they are common (i.e., seagrass beds), and explored mechanistic processes that may drive blooms of this taxa. To establish that human disturbances (e.g., elevated nutrient availability) may drive increased abundance and size of benthic jellyfish, Cassiopea spp., I conducted surveys in human-impacted and unimpacted coastal sites. Jellyfish were more abundant (and larger) from human-impacted areas, positively correlated to elevated nutrient availability. In order to elucidate mechanisms linking Cassiopea spp. with elevated nutrients, I evaluated whether zooxanthellae from Cassiopea were higher from human-disturbed systems, and whether Cassiopea exhibited increased size following nutrient input. I demonstrated that zooxanthellae population densities were elevated in human-impacted sites, and that nutrients led to positive jellyfish growth. As heightened densities of Cassiopea jellyfish may exert top-down and bottom-up controls on flora and fauna in impacted seagrass beds, I sought to examine ecological responses to Cassiopea. I evaluated whether there was a relationship between high Cassiopea densities and lower benthic fauna abundance and diversity in shallow seagrass beds. I found that Cassiopea have subtle effects on benthic fauna. However, through an experiment conducted in a seagrass bed in which nutrients and Cassiopea were added, I demonstrated that Cassiopea can result in seagrass habitat modification, with negative consequences for benthic fauna. My dissertation research demonstrates that increased human-driven benthic jellyfish densities may have indirect and direct effects on flora and fauna of coastal marine systems. This knowledge will advance our understanding of how human disturbances shift species interactions in coastal ecosystems, and will be critical for effective management of jellyfish blooms.

(Table 1) Characteristics of marine snow and ice samples at Barrow, Alaska

We measured light absorption in 42 marine snow, sea ice, seawater, brine, and frost flower samples collected during the OASIS field campaign between February 27 and April 15, 2009. Samples represented multiple sites between landfast ice and open pack ice in coastal areas approximately 5 km west of Barrow, Alaska. The chromophores that are most commonly measured in snow, H2O2, NO3-, and NO2-, on average account for less than 1% of sunlight absorption in our samples. Instead, light absorption is dominated by unidentified "residual" species, likely organic compounds. Light absorption coefficients for the frost flowers on first-year sea ice are, on average, 40 times larger than values for terrestrial snow samples at Barrow, suggesting very large rates of photochemical reactions in frost flowers. For our marine samples the calculated rates of sunlight absorption and OH production from known chromophores are (0.1-1.4) x 10**14 (photons/cm**3/s) and (5-70) x 10**-12 (mol/L/s), respectively. Our residual spectra are similar to spectra of marine chromophoric dissolved organic matter (CDOM), suggesting that CDOM is the dominant chromophore in our samples. Based on our light absorption measurements we estimate dissolved organic carbon (DOC) concentrations in Barrow seawater and frost flowers as approximately 130 and 360 µM C, respectively. We expect that CDOM is a major source of OH in our marine samples, and it is likely to have other significant photochemistry as well.

Hydrogen isotope compositions of terrestrial plant-wax n-alkanes analysed on a transect of marine surface sediment samples from 1°N to 28°S off southwest Africa

A transect of marine surface sediment samples from 1° N to 28° S off southwest Africa was analysed to verify the application of hydrogen isotope compositions of terrestrial plant-wax n-alkanes preserved in ocean sediments as a proxy for continental hydrological conditions. Conditions on the adjacent continent range from humid evergreen forests to deciduous forests, wood- and shrub land and further to arid grasslands and deserts. The hydrogen isotope values for the dominant n-alkane homologues (C29, C31 and C33) vary from -123 per mil to -141 per mil VSMOW and correlate with the modelled hydrogen isotope composition of mean annual and growing season precipitation of postulated continental source areas (r up to 0.8, p < 0.01). The apparent hydrogen isotope fractionation between alkanes and mean annual precipitation is remarkably uniform (-109 per mil on average, Sigma <= 5 per mil, n = 27). Potentially, effects of aridity on the apparent hydrogen isotope fractionation are concealed by the contribution of different plants (C3 dicotyledons vs C4 grasses). Thus, isotope ratios of leaf wax n-alkanes preserved in ocean margin sediments in these and similar tropical regions may be directly converted to dD ratios of ancient precipitation by employing a constant hydrogen isotope fractionation.

n-Alkane content and compound-specific d13C values of soil samples, river samples, and marine surface samples

Southwestern Africa's coastal marine mudbelt, a prominent Holocene sediment package, provides a valuable archive for reconstructing terrestrial palaeoclimates on the adjacent continent. While the origin of terrestrial inorganic material has been intensively studied, the sources of terrigenous organic material deposited in the mudbelt are yet unclear. In this study, plant wax derived n-alkanes and their compound-specific d13C in soils, flood deposits and suspension loads from regional fluvial systems and marine sediments are analysed to characterize the origin of terrestrial organic material in the southwest African mudbelt. Soils from different biomes in the catchments of the Orange River and small west coast rivers show on average distinct n-alkane distributions and compound-specific d13C values reflecting biome-specific vegetation types, most notably the winter rainfall associated Fynbos Biome of the southwestern Cape. In the fluvial sediment samples from the Orange River, changes in the n-alkane distributions and compound-specific d13C compositions reveal an overprint by local vegetation along the river's course. The smaller west coast rivers show distinct signals, reflecting their small catchment areas and particular vegetation communities. Marine surface sediments spanning a transect from the northern mudbelt (29°S) to St. Helena Bay (33°S) reveal subtle, but spatially coherent, changes in n-alkane distributions and compound-specific d13C, indicating the influence of Orange River sediments in the northern mudbelt, the increasing importance of terrigenous input from the adjacent western coastal biomes in the central mudbelt, and contributions from the Fynbos Biome to the southern mudbelt. These findings indicate the different sources of terrestrial organic material deposited in the mudbelt, and highlight the potential the mudbelt has to preserve evidence of environmental change from the adjacent continent.

(Table 3, page 280) Chemical composition of micronodules from a series of stations and sediment depths from Areas C, F, G and K in the Pacific Ocean from R/V Sonne Cruise SO06

Compositional data for coexisting manganese nodules, micronodules, sediments and pore waters from five areas in the equatorial and S.W. Pacific have been obtained. This represents the largest study of its type ever undertaken to establish the distribution of elements between the various phases within the sediment column. The composition of manganese nodules, micronodules and sediments (on a carbonate-free basis) shows marked differences between the equatorial high productivity zone and the low productivity region of the S.W. Pacific. In the case of the nodules, th is reflects an increased supply of transition elements (notably Ni, Cu and Zn) to the nodules as a result of the in situ dissolution of siliceous tests within the sediment column in the equatorial Pacific high productivity zone. Micronodules display similar, but somewhat different, compositions to those of the associated nodules in each area. Micronodule composition is therefore influenced by the same basic factors that control nodule composition, but is modified by dissolution of the micronodules in situ within the sediment column. Locally, as in the area immediately south of the Marquesas Fracture Zone, the micronodule population is contaminated by small, angular volcanic rock fragments; this leads to apparently anomalous micronodule compositions. Micronodules appear to be a transient feature in the sediment column, especially in the equatorial Pacific. Dissolution of micronodules in the sediment column therefore represents an important source of elements for the growth of manganese nodules in the equatorial Pacific. Sediment composition is markedly influenced by the carbonate content. On a carbonate-free basis, the sediments from the equatorial high productivity zone are quite distinct in composition from those in the S.W. Pacific. This reflects differences in the lithology of the sediments. In the Aitutaki Passage, the local influence of volcanoclastic material in sediment composition has been established. The major cations and anions in pore waters measured here show no major differences between equatorial and S.W. Pacific sediments. Silica is, however, higher in equatorial Pacific pore waters reflecting the dissolution of siliceous tests in these sediments.

Benthic freshwater diatom abundance in streams and seepage areas of northern James Ross Island between 2004-2009

Background and aim - The non-marine diatom communities in the Antarctic Region are characterized by a typical species composition, in close relationship with their environment. Despite the growing interest, the diatom flora of James Ross Island is only poorly known. The present paper discusses the diversity of limnoterrestrial diatoms on this island: seepages and streams. Methods - The diatom flora of 53 samples taken on the eastern side of the Ulu peninsula on James Ross Island has been studied using light and scanning electron microscopy. Key results - A total of 69 diatom taxa belonging to 26 genera have been observed. The genera Luticola, Diadesmis, Muelleria and Pinnularia dominated the species composition. The flora shows an interesting mixture of cosmopolitan and Antarctic species containing several species reaching on James Ross Island their most northern distribution in the Antarctic Region. The taxonomical position of one widespread Antarctic species, Psammothidium papilio (D.E.Kellogg, Stuiver, T.B.Kellogg & Denton) Kopalova & Van de Vijver comb. nov., is corrected. Conclusions - The limnoterrestrial diatom flora of James Ross Island has a rather low number of species, of which a large proportion shows a restricted Antarctic distribution.

Description of manganese nodules and crust collected from the Hakuho Maru Cruise KH-69-2, April-June, 1969, in the Japan Trench and Sea of Japan areas

The cores and dredges described in this report were taken on the KH-69-2 Expedition in April-June, 1969 by the Ocean Research Institute, University of Tokyo from the Hakuho Maru. A total of 12 cores and dredges sites have been recovered.

Fossil associations from the middle and upper Eocene strata of the Pamplona Basin and surrounding areas (Navarre, western Pyrenees)

Fossil associations from the middle and upper Eocene (Bartonian and Priabonian) sedimentary succession of the Pamplona Basin are described. This succession was accumulated in the western part of the South Pyrenean peripheral foreland basin and extends from deep-marine turbiditic (Ezkaba Sandstone Formation) to deltaic (Pamplona Marl, Ardanatz Sandstone and Ilundain Marl formations) and marginal marine deposits (Gendulain Formation). The micropalaeontological content is high. It is dominated by foraminifera, and common ostracods and other microfossils are also present. The fossil ichnoasssemblages include at least 23 ichnogenera and 28 ichnospecies indicative of Nereites, Cruziana, Glossifungites and ?Scoyenia-Mermia ichnofacies. Body macrofossils of 78 taxa corresponding to macroforaminifera, sponges, corals, bryozoans, brachiopods, annelids, molluscs, arthropods, echinoderms and vertebrates have been identified. Both the number of ichnotaxa and of species (e. g. bryozoans, molluscs and condrichthyans) may be considerably higher. Body fossil assemblages are comparable to those from the Eocene of the Nord Pyrenean area (Basque Coast), and also to those from the Eocene of the west-central and eastern part of South Pyrenean area (Aragon and Catalonia). At the European scale, the molluscs assemblages seem endemic from the Pyrenean area, although several Tethyan (Italy and Alps) and Northern elements (Paris basin and Normandy) have been recorded. Palaeontological data of studied sedimentary units fit well with the shallowing process that throughout the middle and late Eocene occurs in the area, according to the sedimentological and stratigraphical data.

Application of computer-aided tomography techniques to visualize kelp holdfast structure reveals the importance of habitat complexity for supporting marine biodiversity

Ecosystem engineers that increase habitat complexity are keystone species in marine systems, increasing shelter and niche availability, and therefore biodiversity. For example, kelp holdfasts form intricate structures and host the largest number of organisms in kelp ecosystems. However, methods that quantify 3D habitat complexity have only seldom been used in marine habitats, and never in kelp holdfast communities. This study investigated the role of kelp holdfasts (Laminaria hyperborea) in supporting benthic faunal biodiversity. Computer-aided tomography (CT-) scanning was used to quantify the three-dimensional geometrical complexity of holdfasts, including volume, surface area and surface fractal dimension (FD). Additionally, the number of haptera, number of haptera per unit of volume, and age of kelps were estimated. These measurements were compared to faunal biodiversity and community structure, using partial least-squares regression and multivariate ordination. Holdfast volume explained most of the variance observed in biodiversity indices, however all other complexity measures also strongly contributed to the variance observed. Multivariate ordinations further revealed that surface area and haptera per unit of volume accounted for the patterns observed in faunal community structure. Using 3D image analysis, this study makes a strong contribution to elucidate quantitative mechanisms underlying the observed relationship between biodiversity and habitat complexity. Furthermore, the potential of CT-scanning as an ecological tool is demonstrated, and a methodology for its use in future similar studies is established. Such spatially resolved imager analysis could help identify structurally complex areas as biodiversity hotspots, and may support the prioritization of areas for conservation.

Application of computer-aided tomography techniques to visualize kelp holdfast structure reveals the importance of habitat complexity for supporting marine biodiversity

Ecosystem engineers that increase habitat complexity are keystone species in marine systems, increasing shelter and niche availability, and therefore biodiversity. For example, kelp holdfasts form intricate structures and host the largest number of organisms in kelp ecosystems. However, methods that quantify 3D habitat complexity have only seldom been used in marine habitats, and never in kelp holdfast communities. This study investigated the role of kelp holdfasts (Laminaria hyperborea) in supporting benthic faunal biodiversity. Computer-aided tomography (CT-) scanning was used to quantify the three-dimensional geometrical complexity of holdfasts, including volume, surface area and surface fractal dimension (FD). Additionally, the number of haptera, number of haptera per unit of volume, and age of kelps were estimated. These measurements were compared to faunal biodiversity and community structure, using partial least-squares regression and multivariate ordination. Holdfast volume explained most of the variance observed in biodiversity indices, however all other complexity measures also strongly contributed to the variance observed. Multivariate ordinations further revealed that surface area and haptera per unit of volume accounted for the patterns observed in faunal community structure. Using 3D image analysis, this study makes a strong contribution to elucidate quantitative mechanisms underlying the observed relationship between biodiversity and habitat complexity. Furthermore, the potential of CT-scanning as an ecological tool is demonstrated, and a methodology for its use in future similar studies is established. Such spatially resolved imager analysis could help identify structurally complex areas as biodiversity hotspots, and may support the prioritization of areas for conservation.

Descriptions of Scottish Priority Marine Features (PMFs).

Background The seas around Scotland are rich and diverse – Scotland’s position at the edge of the continental shelf, the long coastline, large area of sea and the mixing of warm and coldwater currents combine to make its waters a special place for marine wildlife and habitats. Scotland has over 18,000 km of coastline and its inshore and offshore areas are among the largest of any EU country, representing 13% of all European seas. Scotland’s seas are of outstanding scenic, historical and cultural value and are part of the national identity at home and abroad. The Marine (Scotland) Act 2010 and the UK Marine and Coastal Access Act 2009 include new powers and duties to ensure that our seas are managed sustainably for future generations, integrating the economic growth of marine industries with the need to protect these assets. Measures to conserve Scotland’s marine natural heritage are based on a three pillar approach, with action at the wider seas level (e.g. marine planning or sectoral controls); specific species conservation measures (e.g. improved protection for seals); and through site protection measures - the identification of new Marine Protected Areas (MPAs). To help target action under each of the three pillars, Scottish Natural Heritage (SNH) and the Joint Nature Conservation Committee (JNCC) have generated a focused list of habitats and species of priority conservation importance - the Priority Marine Features (PMFs). The aim of the current study was to produce a descriptive catalogue of the Scottish PMFs (including component habitats and species where appropriate) to serve as a reference for future nature conservation action. Whilst derived from available existing accounts, the succinct 1-page descriptions are written from a Scottish perspective, refining, but clearly linking to more generic UK, EC or OSPAR (Oslo and Paris Commission) commentary. Available information on the geographic distribution of the features was collated as part of the project and a summary map is provided in each description. Main findings  This project has generated a descriptive catalogue of the 81 PMFs that have been identified in the seas around Scotland (out to the limit of the UK continental shelf). The list comprises 26 broad habitats (e.g. burrowed mud), seven low or limited mobility species (e.g. ocean quahog) and 48 mobile species, including fish (e.g. blue ling) and marine mammals (e.g. minke whale).  Information on the distribution of the PMFs was collated within a Geographic Information System (GIS). This is the first time that data about such a diverse range of Scottish marine nature conservation interests have been compiled within a single repository. These data have and will be used in conjunction with other contextual base-mapping to inform the development of nature conservation advice and commentary (e.g. in the production of the Scotland’s Marine Atlas - Baxter et al., 2011).  The feature distribution mapping used in the production of this report is being made available to view online via the National Marine Plan Interactive web portal (NMPi - http://www.gov.scot/Topics/marine/seamanagement/nmpihome). As new or refined data on Scottish PMFs become available, these will be fed into updates to the project geodatabase and NMPi.

Descriptions of Scottish Priority Marine Features (PMFs).

Background The seas around Scotland are rich and diverse – Scotland’s position at the edge of the continental shelf, the long coastline, large area of sea and the mixing of warm and coldwater currents combine to make its waters a special place for marine wildlife and habitats. Scotland has over 18,000 km of coastline and its inshore and offshore areas are among the largest of any EU country, representing 13% of all European seas. Scotland’s seas are of outstanding scenic, historical and cultural value and are part of the national identity at home and abroad. The Marine (Scotland) Act 2010 and the UK Marine and Coastal Access Act 2009 include new powers and duties to ensure that our seas are managed sustainably for future generations, integrating the economic growth of marine industries with the need to protect these assets. Measures to conserve Scotland’s marine natural heritage are based on a three pillar approach, with action at the wider seas level (e.g. marine planning or sectoral controls); specific species conservation measures (e.g. improved protection for seals); and through site protection measures - the identification of new Marine Protected Areas (MPAs). To help target action under each of the three pillars, Scottish Natural Heritage (SNH) and the Joint Nature Conservation Committee (JNCC) have generated a focused list of habitats and species of priority conservation importance - the Priority Marine Features (PMFs). The aim of the current study was to produce a descriptive catalogue of the Scottish PMFs (including component habitats and species where appropriate) to serve as a reference for future nature conservation action. Whilst derived from available existing accounts, the succinct 1-page descriptions are written from a Scottish perspective, refining, but clearly linking to more generic UK, EC or OSPAR (Oslo and Paris Commission) commentary. Available information on the geographic distribution of the features was collated as part of the project and a summary map is provided in each description. Main findings  This project has generated a descriptive catalogue of the 81 PMFs that have been identified in the seas around Scotland (out to the limit of the UK continental shelf). The list comprises 26 broad habitats (e.g. burrowed mud), seven low or limited mobility species (e.g. ocean quahog) and 48 mobile species, including fish (e.g. blue ling) and marine mammals (e.g. minke whale).  Information on the distribution of the PMFs was collated within a Geographic Information System (GIS). This is the first time that data about such a diverse range of Scottish marine nature conservation interests have been compiled within a single repository. These data have and will be used in conjunction with other contextual base-mapping to inform the development of nature conservation advice and commentary (e.g. in the production of the Scotland’s Marine Atlas - Baxter et al., 2011).  The feature distribution mapping used in the production of this report is being made available to view online via the National Marine Plan Interactive web portal (NMPi - http://www.gov.scot/Topics/marine/seamanagement/nmpihome). As new or refined data on Scottish PMFs become available, these will be fed into updates to the project geodatabase and NMPi.