Biogeographic Patterns, Predator Identity, and Chemical Signals Influence the Occurrence and Magnitude of Non-lethal Predator Effects

Predators can have large effects on prey populations and on the structure and function of communities. In addition to direct consumption of prey, predators often cause prey to alter their foraging behavior, habitat selection, and morphology. These non-lethal effects of predators can propagate to multiple trophic levels and often exert equal or larger effects upon communities than those of direct consumption. For non-lethal predatory effects to occur, prey must detect and respond to predation risk. While the importance of information transfer in this process has been realized, few studies explore how prey responses are influenced by predator characteristics and environmental conditions that influence the transmission of cues indicative of predation risk. In this dissertation I investigate factors that influence how a single prey species evaluates and responds to predation risk. Here, I examined: 1) the type and nature of cues prey use to evaluate predator risk; 2) how predator identity, predator diet, and the relative risk of predators influence prey response to predation risk; 3) how hydrodynamic conditions influence the delivery of predator cues; 4) how biogeographic trends in predator distribution influence prey response to predation risk; and 5) how genetic structure might vary according to prey geographic location and habitat. To address these questions, I used a common intertidal model system consisting of the rocky intertidal whelk Nucella lapillus (Linnaeus, 1758) and a suite of its predators, the native rock crab Cancer irroratus (Say, 1817), Jonah crab Cancer borealis (Stimpson, 1859), and the invasive green crab Carcinus maenas (Linnaeus, 1758). Nucella use chemical cues emanating from their most common predator (Carcinus maenas) and crushed conspecifics to evaluate predation risk. Nucella from different habitats experience different levels of predation risk, and Nucella from habitats with high levels of predation had larger antipredatory responses to predator risk cues than Nucella that experienced less predation. These chemical cues indicative of predation risk are influenced by hydrodynamic conditions, and Nucella have the strongest anti-predatory response in flow velocities of u= ~4- 8 cm s^-1. Furthermore, Nucella from geographic regions where green crabs are historically absent did not elicit anti-predatory responses, while Nucella from regions where green crabs are common frequently responded. Findings from my dissertation research demonstrate that prey detection and response to predation risk is highly dependent upon predator identity, predator diet, environmental forces, and biogeographic patterns in predator and prey distributions

Combined sulfur, carbon and redox budget constraints on genetic models for the Here's Your Chance Pb-Zn deposit, Australia

The formation of base metal sulfide deposits requires not only a source of metals but also reduced sulfur. If incoming sulfur is present in ore fluids as sulfate, then a source of electrons is needed to drive the reduction of sulfate to sulfide. The oxidation of organic matter can release electrons that provide the reducing capacity, whether it be in low- or high-temperature settings that are conducive to biological or thermochemical sulfate reduction (BSR or TSR). The amounts of organic matter reacted and sulfide minerals formed can be estimated by mass balance calculations. In this study, an integrated mass balance expression is formulated that takes into account the sulfide mineral content and organic carbon content and H/C ratios of mineralised and non-mineralised rocks. Model calculations based on carbon, sulfur and redox budget balances suggest that the extent of oxidation of the organic matter present at the Here’s Your Chance (HYC) Pb–Zn deposit is insufficient for reduction of the required quantity of sulfate. The results imply that externally derived reducing capacity and/or reduced sulfur is required to form the metal resource. Possible sources include hydrocarbon-rich fluids from deeper parts of the sedimentary sequence or formation of sulfide and organic matter as products of BSR during sedimentation/early diagenesis. However, the observed oxidation of organic matter associated with the deposit suggests that at least some reducing capacity is locally derived. Therefore, our calculations are consistent with genetic models for HYC that have multiple sources of redox budget for sulfate reduction

Neural Responses to Complex Auditory Rhythms: The Role of Attending

The aim of this study was to explore the role of attention in pulse and meter perception using complex rhythms. We used a selective attention paradigm in which participants attended to either a complex auditory rhythm or a visually presented word list. Performance on a reproduction task was used to gauge whether participants were attending to the appropriate stimulus. We hypothesized that attention to complex rhythms – which contain no energy at the pulse frequency – would lead to activations in motor areas involved in pulse perception. Moreover, because multiple repetitions of a complex rhythm are needed to perceive a pulse, activations in pulse-related areas would be seen only after sufficient time had elapsed for pulse perception to develop. Selective attention was also expected to modulate activity in sensory areas specific to the modality. We found that selective attention to rhythms led to increased BOLD responses in basal ganglia, and basal ganglia activity was observed only after the rhythms had cycled enough times for a stable pulse percept to develop. These observations suggest that attention is needed to recruit motor activations associated with the perception of pulse in complex rhythms. Moreover, attention to the auditory stimulus enhanced activity in an attentional sensory network including primary auditory cortex, insula, anterior cingulate, and prefrontal cortex, and suppressed activity in sensory areas associated with attending to the visual stimulus

Mechanisms governing interannual variability of upper-ocean temperature in a global ocean hindcast simulation

Author Posting. © American Meteorological Society, 2007. This article is posted here by permission of American Meteorological Society for personal use, not for redistribution. The definitive version was published in Journal of Physical Oceanography 37 (2007): 1918-1938, doi:10.1175/jpo3089.1.The interannual variability in upper-ocean (0–400 m) temperature and governing mechanisms for the period 1968–97 are quantified from a global ocean hindcast simulation driven by atmospheric reanalysis and satellite data products. The unconstrained simulation exhibits considerable skill in replicating the observed interannual variability in vertically integrated heat content estimated from hydrographic data and monthly satellite sea surface temperature and sea surface height data. Globally, the most significant interannual variability modes arise from El Niño–Southern Oscillation and the Indian Ocean zonal mode, with substantial extension beyond the Tropics into the midlatitudes. In the well-stratified Tropics and subtropics, net annual heat storage variability is driven predominately by the convergence of the advective heat transport, mostly reflecting velocity anomalies times the mean temperature field. Vertical velocity variability is caused by remote wind forcing, and subsurface temperature anomalies are governed mostly by isopycnal displacements (heave). The dynamics at mid- to high latitudes are qualitatively different and vary regionally. Interannual temperature variability is more coherent with depth because of deep winter mixing and variations in western boundary currents and the Antarctic Circumpolar Current that span the upper thermocline. Net annual heat storage variability is forced by a mixture of local air–sea heat fluxes and the convergence of the advective heat transport, the latter resulting from both velocity and temperature anomalies. Also, density-compensated temperature changes on isopycnal surfaces (spice) are quantitatively significant.This work was supported in part from NOAA Office of Global Programs ACCP Grant NA86GP0290, NSF Grant OCE96-33681, and the WHOI Ocean and Climate Change Institute

Impact of eddy–wind interaction on eddy demographics and phytoplankton community structure in a model of the North Atlantic Ocean

Author Posting. © The Author(s), 2010. This is the author's version of the work. It is posted here by permission of Elsevier B.V. for personal use, not for redistribution. The definitive version was published in Dynamics of Atmospheres and Oceans 52 (2011): 80-94, doi:10.1016/j.dynatmoce.2011.01.003.Two eddy-resolving (0.1-degree) physical-biological simulations of the North Atlantic Ocean are compared, one with the surface momentum flux computed only from wind velocities and the other using the difference between air and ocean velocity vectors. This difference in forcing has a significant impact on the intensities and relative number of different types of mesoscale eddies in the Sargasso Sea. Eddy/wind interaction significantly reduces eddy intensities and increases the number of mode-water eddies and “thinnies” relative to regular cyclones and anticyclones; it also modifies upward isopycnal displacements at the base of the euphotic zone, increasing them in the centers of mode water eddies and at the edges of cyclones, and decreasing them in the centers of cyclones. These physical changes increase phytoplankton growth rates and biomass in mode-water eddies, bringing the biological simulation into better agreement with field data. These results indicate the importance of including the eddy/wind interaction in simulations of the physics and biology of eddies in the subtropical North Atlantic. However, eddy intensities in the simulation with eddy/wind interaction are lower than observed, which suggests a decrease in horizontal viscosity or an increase in horizontal grid resolution will be necessary to regain the observed level of eddy activity.LAA and DJM gratefully acknowledge the support of NASA grant 07-CARBON07-17. SCD and IDL gratefully acknowledge support from the NSF Center for Microbial Oceanography, Research and Education (C-MORE; NSF EF-0424599)

Spring bloom dynamics and zooplankton biomass response on the US Northeast Continental Shelf

This paper is not subject to U.S. copyright. The definitive version was published in Continental Shelf Research 102 (2015): 47-61, doi:10.1016/j.csr.2015.04.005.The spring phytoplankton bloom on the US Northeast Continental Shelf is a feature of the ecosystem production cycle that varies annually in timing, spatial extent, and magnitude. To quantify this variability, we analyzed remotely-sensed ocean color data at two spatial scales, one based on ecologically defined sub-units of the ecosystem (production units) and the other on a regular grid (0.5°). Five units were defined: Gulf of Maine East and West, Georges Bank, and Middle Atlantic Bight North and South. The units averaged 47×103 km2 in size. The initiation and termination of the spring bloom were determined using change-point analysis with constraints on what was identified as a bloom based on climatological bloom patterns. A discrete spring bloom was detected in most years over much of the western Gulf of Maine production unit. However, bloom frequency declined in the eastern Gulf of Maine and transitioned to frequencies as low as 50% along the southern flank of the Georges Bank production unit. Detectable spring blooms were episodic in the Middle Atlantic Bight production units. In the western Gulf of Maine, bloom duration was inversely related to bloom start day; thus, early blooms tended to be longer lasting and larger magnitude blooms. We view this as a phenological mismatch between bloom timing and the “top-down” grazing pressure that terminates a bloom. Estimates of secondary production were available from plankton surveys that provided spring indices of zooplankton biovolume. Winter chlorophyll biomass had little effect on spring zooplankton biovolume, whereas spring chlorophyll biomass had mixed effects on biovolume. There was evidence of a “bottom up” response seen on Georges Bank where spring zooplankton biovolume was positively correlated with the concentration of chlorophyll. However, in the western Gulf of Maine, biovolume was uncorrelated with chlorophyll concentration, but was positively correlated with bloom start and negatively correlated with magnitude. This observation is consistent with both a “top-down” mechanism of control of the bloom and a “bottom-up” effect of bloom timing on zooplankton grazing. Our inability to form a consistent model of these relationships across adjacent systems underscores the need for further research

Assessment of hydropyrolysis as a method for the quantification of black carbon using standard reference materials

A wide selection of thermal, chemical and optical methods have been proposed for the quantification of black carbon (BC) in environmental matrices, and the results to date differ markedly depending upon the method used. A new approach is hydropyrolysis (hypy), where pyrolysis assisted by high hydrogen pressures (150 bar) facilitates the complete reductive removal of labile organic matter, so isolating a highly stable portion of the BC continuum (defined as BChypy). Here, the potential of hypy for the isolation and quantification of BC is evaluated using the 12 reference materials from the International BC Ring Trial, comprising BC-rich samples, BC-containing environmental matrices and BC-free potentially interfering materials. By varying the hypy operating conditions, it is demonstrated that lignocellulosic, humic and other labile organic carbon material (defined as non-BChypy) is fully removed by 550 °C, with hydrogasification of the remaining BChypy not commencing until over 575 °C. The resulting plateau in sample mass and carbon loss is apparent in all of the environmental samples, facilitating BC quantification in a wide range of materials. The BChypy contents for all 12 ring trial samples fall within the range reported in the BC inter-comparison study, and systematic differences with other methods are rationalised. All methods for BC isolation, including hypy are limited by the fact that BC cannot be distinguished from extremely thermally mature organic matter; for example in high rank coals. However, the data reported here indicates that BChypy has an atomic H/C ratio of less than 0.5 and therefore comprises a chemically well-defined polyaromatic structure in terms of the average size of peri-condensed aromatic clusters of >7 rings (24 carbon atoms), that is consistent across different sample matrices. This, together with the sound underlying rationale for the reductive removal of labile organic matter, makes hypy an ideal approach for matrix independent BC quantification. The hypy results are extremely reproducible, with BChypy determinations from triplicate analyses typically within ±2% across all samples, limited mainly by the precision of the elemental analyser

Towards a free-free template for CMB foregrounds

A full-sky template map of the Galactic free-free foreground emission component is increasingly important for high sensitivity CMB experiments. We use the recently published \ha data of both the northern and southern skies as the basis for such a template. The first step is to correct the \ha maps for dust absorption using the 100 $\mu$m dust maps of Schlegel, Finkbeiner & Davis (1998). We show that for a range of longitudes, the Galactic latitude distribution of absorption suggests that it is 33 per cent of the full extragalactic absorption. A reliable absorption-corrected \ha map can be produced for $\sim 95$ per cent of the sky; the area for which a template cannot be recovered is the Galactic plane area $|b| < 5^{\circ}$, $l=260^{\circ}-0^{\circ}-160^{\circ}$ and some isolated dense dust clouds at intermediate latitudes. The second step is to convert the dust-corrected \ha data into a predicted radio surface brightness. The free-free emission formula is revised to give an accurate expression (1 per cent) for the radio emission covering the frequency range 100 MHz to 100 GHz and the electron temperature range 3000 to 20000 K. The main uncertainty when applying this expression is the variation of electron temperature across the sky. The emission formula is verified in several extended H{\sc ii} regions using data in the range 408 to 2326 MHz. A full-sky free-free template map is presented at 30 GHz; the scaling to other frequencies is given. The Haslam et al. all-sky 408 MHz map of the sky can be corrected for this free-free component, which amounts to a $\approx 6$ per cent correction at intermediate and high latitudes....Comment: 18 pages, 11 figures, accepted for publication in M.N.R.A.S. High-resolution versions of figs 2,7 (in colour), 9 and 11 can be obtained from ftp://ftp.jb.man.ac.uk/pub/cdickins/ff_paper/FINAL_FIGURES

Ocean chlorofluorocarbon and heat uptake during the twentieth century in the CCSM3

Author Posting. © American Meteorological Society 2006. This article is posted here by permission of American Meteorological Society for personal use, not for redistribution. The definitive version was published in Journal of Climate 19 (2006): 2366–2381, doi:10.1175/JCLI3758.1.An ensemble of nine simulations for the climate of the twentieth century has been run using the Community Climate System Model version 3 (CCSM3). Three of these runs also simulate the uptake of chlorofluorocarbon-11 (CFC-11) into the ocean using the protocol from the Ocean Carbon Model Intercomparison Project (OCMIP). Comparison with ocean observations taken between 1980 and 2000 shows that the global CFC-11 uptake is simulated very well. However, there are regional biases, and these are used to identify where too much deep-water formation is occurring in the CCSM3. The differences between the three runs simulating CFC-11 uptake are also briefly documented. The variability in ocean heat content in the 1870 control runs is shown to be only a little smaller than estimates using ocean observations. The ocean heat uptake between 1957 and 1996 in the ensemble is compared to the recent observational estimates of the secular trend. The trend in ocean heat uptake is considerably larger than the natural variability in the 1870 control runs. The heat uptake down to 300 m between 1957 and 1996 varies by a factor of 2 across the ensemble. Some possible reasons for this large spread are discussed. There is much less spread in the heat uptake down to 3 km. On average, the CCSM3 twentieth-century ensemble runs take up 25% more heat than the recent estimate from ocean observations. Possible explanations for this are that the model heat uptake is calculated over the whole ocean, and not just in the regions where there are many observations and that there is no parameterization of the indirect effects of aerosols in CCSM3.Support provided by the National Science Foundation, the Department of Energy, the Ministry of Education, Culture, Sports, Science and Technology, and the Earth Simulator Center of the Japan Agency for Marine- Earth Science and Technology