Functional responses for zooplankton feeding on multiple resources: a review of assumptions and biological dynamics

Modelers often need to quantify the rates at which zooplankton consume a variety of species, size classes and trophic types. Implicit in the equations used to describe the multiple resource functional response (i.e. how nutritional intake varies with resource densities) are assumptions that are not often stated, let alone tested. This is problematic because models are sensitive to the details of these formulations. Here, we enable modelers to make more informed decisions by providing them with a new framework for considering zooplankton feeding on multiple resources. We define a new classification of multiple resource responses that is based on preference, selection and switching, and we develop a set of mathematical diagnostics that elucidate model assumptions. We use these tools to evaluate the assumptions and biological dynamics inherent in published multiple resource responses. These models are shown to simulate different resource preferences, implied single resource responses, changes in intake with changing resource densities, nutritional benefits of generalism, and nutritional costs of selection. Certain formulations are further shown to exhibit anomalous dynamics such as negative switching and sub-optimal feeding. Such varied responses can have vastly different ecological consequences for both zooplankton and their resources; inappropriate choices may incorrectly quantify biologically-mediated fluxes and predict spurious dynamics. We discuss how our classes and diagnostics can help constrain parameters, interpret behaviors, and identify limitations to a formulation's applicability for both regional (e.g. High-Nitrate-Low-Chlorophyll regions comprising large areas of the Pacific) and large-scale applications (e.g. global biogeochemical or climate change models). Strategies for assessing uncertainty and for using the mathematics to guide future experimental investigations are also discussed.     

Geomorphological evidence for upslope canyon-forming processes on the northern KwaZulu-Natal shelf,SW Indian Ocean,South Africa

A geomorphological and statistical analysis of slope canyons from the northern KwaZulu-Natal continental margin is documented and compared with submarine canyons from the Atlantic margin of the USA. The northern KwaZulu-Natal margin is characterized by increasing upslope relief, concave slope-gradient profiles and features related to upslope growth of the canyon forms. Discounting slope-gradient profile, this morphology is strikingly similar to canyon systems of the New Jersey slope. Several phases of canyon incision indicate that downslope erosion is also an important factor in the evolution of the northern KwaZulu-Natal canyon systems. Despite the strong similarities between the northern KwaZulu-Natal and New Jersey slope-canyon systems, key differences are evident: (1) the concavity of the northern KwaZulu-Natal slope, contrasting with the ∼linear New Jersey slope; (2) the relative isolation of the northern KwaZulu-Natal canyons, rather than the dense clustering of the New Jersey canyons; and (3) the absence of strongly shelf-breaching canyons along the northern KwaZulu-Natal margin. In comparison with the New Jersey margin, we surmise a more youthful stage of canyon evolution, a result of either the canyons themselves being younger or the formative processes being less active. Less complicated patterns of erosion resulting from reduced sediment availability have developed in northern KwaZulu-Natal. The reduction in slope concavity on the New Jersey margin may be the result of grading of the upper slope by intensive headward erosion, a process more subdued—or less evident—on the KwaZulu-Natal margin.     

An evaluation of metal bioaccessibility in estuarine sediments using the commercially available protein, bovine serum albumin

The bioaccessibility of metals (Al, Ca, Fe, Mn, Ag, Cd, Co, Cu, Ni, Pb, Sn, Zn) in oxic estuarine sediments has been evaluated using solutions of a commercially available protein (bovine serum albumin; BSA) that mimic the chemical conditions encountered in the gut environment of many deposit-feeding organisms. Over a 20 h incubation period with 5 g L^− 1 BSA, metal mobilisation was generally biphasic in that a relatively short period of rapid release was succeeded by more gradual release or approach to equilibrium, although in some cases metal readsorption was evident during the time-courses. Availability to BSA, defined as metal released after 20 h relative to metal extracted by boiling aqua regia, was greatest for Cd, Ni, Ca and Zn and lowest for Fe and Mn, and correlated well with, but was an order of magnitude lower than, metal digested by acidified hydrogen peroxide. Time-courses conducted in the absence of the protein revealed that significant quantities of Ca and Mn were water-soluble, reflecting the partial dissolution of carbonates and hydrous Mn oxides, respectively. Additional experiments indicated a net increase in metal release with increasing BSA concentration and, with the exception of Ca and Mn, a substantial increase in metal mobilisation after sediment organic matter had been digested by peroxidation. These observations suggest that, apart from Ca and Mn, metal release proceeds via complexation with component amino acids of the protein, denudation of organic host phases, and exposure of inorganic, metal-bearing minerals. Accordingly, the bioaccessibility of a metal is predicted to be dependent on its (i) affinity for proteinaceous ligands, (ii) association with components of digestible sediment organic matter, and (iii) degree of binding at relatively weak sites on sediment phases that are exposed or modified by the action of proteins and other chemical constituents of the gut environment.     

Using existing scientific capacity to set targets for ecosystem-based management: A Puget Sound case study

Human actions have altered the structure and function of coastal ecosystems worldwide. In many locations, the overall portfolio of goods, cultural amenities, and supporting services provided by the marine environment has deteriorated. Ecosystem-based management (EBM) offers significant promise for addressing these issues because it is a comprehensive and integrated approach designed to reconcile conflicts and trade-offs among users of marine resources. A key step in the implementation of EBM is the establishment of target reference levels, or desired states, for indicators that reflect the status of the ecosystem. This paper reviews five approaches, borrowed from a variety of disciplines, to establish target reference levels for EBM. The approaches include the use of existing reference levels, reference directions, and reference levels based on nonlinear functional relationships, baselines, or social norms. Each approach is particularly suitable for EBM because it can be used alone or in combination with others to contextualize status for a diverse suite of ecosystem goals influenced by a wide variety of human activities. Perhaps most importantly, these approaches offer a prospectus for moving forward with EBM by using readily available information, motivating existing scientific capacity, and addressing trade-offs implicit to the setting of targets. This last point is articulated via examples of how each type of reference level might be applied in Puget Sound, WA, USA, where the efforts of scientists, managers, and policy makers have aligned recently in the interest of EBM implementation.     

Long lifetimes of β-glucosidase,leucine aminopeptidase,and phosphatase in Arctic seawater

The active lifetime of extracellular enzymes is a critical determinant of the effectiveness of enzyme production as a means for heterotrophic marine microbes to obtain organic substrates. Here, we report lifetimes of three classes of extracellular enzyme in Arctic seawater. We also investigated the relative importance of photochemical processes and particle-associated processes in inactivating extracellular enzymes. Enzyme inactivation in filtered seawater was slow, with apparent half-lives of enzyme activities on the order of hundreds of hours. The presence of particles (including cells) did not significantly change inactivation rates, suggesting that the long half-lives observed in filtered seawater were realistic for enzymes in unfiltered seawater. Phosphatase and leucine aminopeptidase were susceptible to photoinactivation, but only under high intensity UV-B and UV-C illumination; there was no evidence for increased inactivation rates under natural illumination at our study site in Ny Ålesund, Svalbard. Comparison of inactivation rates of commercially-obtained enzymes from non-marine sources with the extracellular enzymes naturally present in Arctic seawater suggests that the natural enzymes contain structural features that confer longer lifetimes, consistent with observations reported by others from a range of field sites that cell-free enzymes can contribute a substantial fraction of total hydrolytic activity in the water column.     

The Regional Ocean Modeling System (ROMS) 4-dimensional variational data assimilation systems: Part II - Performance and application to the California Current System

The Regional Ocean Modeling System (ROMS) 4-dimensional variational (4D-Var) data assimilation systems have been systematically applied to the mesoscale circulation environment of the California Current to demonstrate the performance and practical utility of the various components of ROMS 4D-Var. In particular, we present a comparison of three approaches to 4D-Var, namely: the primal formulation of the incremental strong constraint approach; the dual formulation “physical-space statistical analysis system”; and the dual formulation indirect representer approach. In agreement with theoretical considerations all three approaches converge to the same ocean circulation estimate when using the same observations and prior information. However, the rate of convergence of the dual formulation was found to be inferior to that of the primal formulation. Other aspects of the 4D-Var performance that relate to the use of multiple outer-loops, preconditioning, and the weak constraint are also explored. A systematic evaluation of the impact of the various components of the 4D-Var control vector (i.e. the initial conditions, surface forcing and open boundary conditions) is also presented. It is shown that correcting for uncertainties in the model initial conditions exerts the largest influence on the ability of the model to fit the available observations. Various important diagnostics of 4D-Var are also examined, including estimates of the posterior error, the information content of the observation array, and innovation-based consistency checks on the prior error assumptions. Using these diagnostic tools, we find that more than 90% of the observations assimilated into the model provide redundant information. This is a symptom of the large percentage of satellite data that are used and to some extent the nature of the data processing employed. This is the second in a series of three papers describing the ROMS 4D-Var systems.     

People helping turtles, turtles helping people: Understanding resident attitudes towards sea turtle conservation and opportunities for enhanced community participation in Bahia Magdalena, Mexico

In Pacific Mexico, all five sea turtle species have declined over the past century due to intense overexploitation of meat and eggs, fisheries bycatch, and degradation of marine and nesting habitats. One of the most heavily impacted areas has been the Baja California peninsula, where sea turtle populations remain historically low despite existing conservation measures that include a complete moratorium on the use of sea turtles, over three decades of widespread protection of nesting beaches, and in-water monitoring of sea turtles at coastal foraging areas. We recognize the need for alternative sea turtle conservation strategies that rely on increased participation of civil society and Mexican citizens. The purpose of this paper was to identify resident attitudes towards sea turtle conservation and opportunities for enhanced community participation in Bahia Magdalena, a region in Baja California Sur, Mexico experiencing high levels of sea turtle poaching and bycatch in fisheries. Through semi-structured interviews we found that while residents were overwhelmingly interested in participating in sea turtle conservation, peer pressure and conflict within the community presented major challenges. The majority of residents indicated that sea turtle voluntourism would have a positive impact on their community. Economic incentives and increased protection for sea turtles were mentioned as benefits of sea turtle voluntourism, whereas peer pressure, difficulty obtaining permits and producing effective marketing materials, and doubt about direct economic benefits were cited as constraints. We discuss our results in terms of opportunities, challenges, and recommendations for improving community-focused sea turtle conservation throughout the region.     

First observations of jelly-falls at the seafloor in a deep-sea fjord

Faunal communities at the deep-sea floor mainly rely on the downward transport of particulate organic material for energy, which can come in many forms, ranging from phytodetritus to whale carcasses. Recently, studies have shown that the deep-sea floor may also be subsidized by fluxes of gelatinous material to the benthos. The deep-sea scyphozoan medusa Periphylla periphylla is common in many deep-sea fjords in Norway and recent investigations in Lurefjorden in western Norway suggest that the biomass of this jellyfish currently exceeds 50000 t here. To quantify the presence of dead P. periphylla jellyfish falls (hereafter termed jelly-falls) at the deep seafloor and the standing stock of carbon (C) and nitrogen (N) deposited on the seafloor by this species, we made photographic transects of the seafloor, using a ‘Yo-Yo’ camera system during an opportunistic sampling campaign in March 2011. Of 218 seafloor photographs taken, jelly-falls were present in five, which resulted in a total jelly-fall abundance of 1×10^-2 jelly-falls m⁻² over the entire area surveyed. Summed over the entire area of seafloor photographed, 1×10^-2 jelly-falls m⁻² was equivalent to a C- and N-biomass of 13 mg C m⁻² and 2 mg N m⁻². The contribution of each jelly-fall to the C- and N-amount of the sediment in the immediate vicinity of each fall (i.e. to sediment in each 3.02 m² image in which jelly-falls were observed) was estimated to be 568±84 mg C m⁻² and 88±13 mg N m⁻². The only megafaunal taxon observed around or on top of the jelly-falls was caridean shrimp (14±5 individuals jelly-fall⁻¹), and shrimp abundance was significantly greater in photographs in which a jelly-fall was found (14±5 individuals image⁻¹) compared to photographs in which no jelly-falls were observed (1.4±0.7 individuals image⁻¹). These observations indicate that jelly-falls in this fjord can enhance the sedimentary C- and N-amount at the deep-sea floor and may provide nutrition to benthic and demersal faunas in this environment. However, organic enrichment from the jelly-falls found in this single sampling event and associated disturbance was highly localized.     

Autochthonous inheritance of zircon through Cretaceous partial melting of Carboniferous plutons: the Arthur River Complex,Fiordland, New Zealand

TIMS and SHRIMP U–Pb analyses of zircons from Milford Orthogneiss metadiorite (P = 1–1.4 GPa; T ≥ 750°C) of the Arthur River Complex of northern Fiordland reveal a bimodal age pattern. Zircons are predominantly either Paleozoic (357.0 ± 4.2 Ma) and prismatic with oscillatory zoning, or Cretaceous (133.9 ± 1.8 Ma) and ovoid with sector or patchy zoning. The younger age component is not observed overgrowing older grains. Most grains of both ages are overgrown by younger Cretaceous (~120 Ma) metamorphic zircon with very low U and Th/U (0.01). We interpret the bimodal ages as indicating initial igneous emplacement and crystallisation of a dioritic protolith pluton at ~357 Ma, followed by Early Cretaceous granulite-facies metamorphism at ~134 Ma, during which a significant fraction (~60%) of the zircon grains dissolved, and subsequently reprecipitated, effectively in situ, in partial melt pockets. The remaining ~40% of original Paleozoic grains were apparently not in contact with the partial melt, remained intact, and show only slight degrees of Pb loss. Sector zoning of the Cretaceous grains discounts their origin by solid state recrystallisation of Paleozoic grains. The alternative explanation—that the Paleozoic component represents a 40% inherited component in an Early Cretaceous transgressive dioritic magma—is considered less likely given the relatively high solubility of zircon in magma of this composition, the absence of 134 Ma overgrowths, the single discrete age of the older component, equivalent time-integrated ¹⁷⁷Hf/¹⁷⁶Hf compositions of both age groups, and the absence of the Cambrian-Proterozoic detrital zircon that dominates regional Cambro-Ordovician metasedimentary populations. Similar bimodal Carboniferous-Early Cretaceous age distributions are characteristic of the wider Arthur River Complex; 8 of 12 previously dated dioritic samples have a Paleozoic component averaging 51%. Furthermore, the age and chemical suite affinity of these and several more felsic rocks can be matched with those of the relatively unmetamorphosed Carboniferous plutonic terrane along the strike of the Mesozoic margin in southern Fiordland, also supporting the in situ derivation of the Carboniferous “inherited” component.