Biodiversity in agricultural landscapes can be increased with conversion of some production lands into 'more-natural'- unmanaged or extensively managed - lands. However, it remains unknown to what extent biodiversity can be enhanced by altering landscape pattern without reducing agricultural production. We propose a framework for this problem, considering separately compositional heterogeneity (the number and proportions of different cover types) and configurational heterogeneity (the spatial arrangement of cover types). Cover type classification and mapping is based on species requirements, such as feeding and nesting, resulting in measures of 'functional landscape heterogeneity'. We then identify three important questions: does biodiversity increase with (1) increasing heterogeneity of the more-natural areas, (2) increasing compositional heterogeneity of production cover types and (3) increasing configurational heterogeneity of production cover types? We discuss approaches for addressing these questions. Such studies should have high priority because biodiversity protection globally depends increasingly on maintaining biodiversity in human-dominated landscapes.

[1] The first quasi-annual time series of nutrients and chlorophyll fluorescence in the southeast Beaufort Sea showed that mixing, whether driven by wind, local convection, or brine rejection, and the ensuing replenishment of nutrients at the surface were minimal during autumn and winter. Anomalously high inventories of nutrients were observed briefly in late December, coinciding with the passage of an eddy generated offshore. The concentrations of NO 3 À in the upper mixed layer were otherwise low and increased slowly from January to April. The coincident decline of NO 2 À suggested nitrification near the surface. The vernal drawdown of NO 3 À in 2004 began at the ice-water interface during May, leaving as little as 0.9 mM of NO 3 À when the ice broke up. A subsurface chlorophyll maximum (SCM) developed promptly and deepened with the nitracline until early August. The diatom-dominated SCM possibly mediated half of the seasonal NO 3 À consumption while generating the primary NO 2 À maximum. Dissolved inorganic carbon and soluble reactive phosphorus above the SCM continued to decline after NO 3 À was depleted, indicating that net community production (NCP) exceeded NO 3 À -based new production. These dynamics contrast with those of productive Arctic waters where nutrient replenishment in the upper euphotic zone is extensive and NCP is fueled primarily by allochthonous NO 3 À . The projected increase in the supply of heat and freshwater to the Arctic should bolster vertical stability, further reduce NO 3 À -based new production, and increase the relative contribution of the SCM. This trend might be reversed locally or regionally by the physical forcing events that episodically deliver nutrients to the upper euphotic zone.

Comprehensive investigations of the Canadian Arctic during late summer and early fall revealed the widespread occurrence of long-lived subsurface chlorophyll maxima (SCM) in seasonally ice-free waters. The vertical position of the SCM corresponded with the depth of the subsurface biomass maximum (SBM), at least in Baffin Bay, suggesting that SCM could be an important source of carbon for the food web. Most of these SCM were located well below the pycnocline in close association with the nitracline, implying that their vertical position was driven mainly by a shortage of inorganic nitrogen in the upper euphotic zone. The diversity of SCM configurations with respect to physical properties of the water column complicates the estimation of euphotic-zone chlorophyll and primary production from surface properties. High photosynthetic yields (F v /F m ) showed the phytoplankton to be photosynthetically competent and well acclimated to conditions of irradiance and nutrient supply near the surface and at the SCM. A well-defined primary nitrite maximum was associated with the SCM in the southwest Canadian Arctic, but not in the northeast where nitrite concentrations were highest much below the euphotic zone. This contrast is consistent with differences in vertical stratification, the light -dark cycle and, possibly, the physiological state and taxonomic composition of the phytoplankton community at the SCM. This study demonstrates that the SCM, once regarded as anecdotal due to under-sampling, are a dominant feature of the Arctic Ocean that should be considered in remote sensing studies and biogeochemical models.

The Canadian Beaufort Sea has been categorized as an oligotrophic system with the potential for enhanced production due to a nutrient‐rich intermediate layer of Pacific‐origin waters. Using under‐ice hydrographic data collected near the ice‐edge of a shallow Arctic bay, we documented an ice‐edge upwelling event that brought nutrient‐rich waters to the surface during June 2008. The event resulted in a 3‐week long phytoplankton bloom that produced an estimated 31 g C m−2 of new production. This value was approximately twice that of previous estimates for annual production in the region, demonstrating the importance of ice‐edge upwelling to the local marine ecosystem. Under‐ice primary production estimates of up to 0.31 g C m−2 d−1 showed that this production was not negligible, contributing up to 22% of the daily averaged production of the ice‐edge bloom. It is suggested that under‐ice blooms are a widespread yet under‐documented phenomenon in polar regions, which could increase in importance with the Arctic's thinning ice cover and subsequent increase in transmitted irradiance to the under‐ice environment.

Quantifying the costs and benefits of migration distance is critical to understanding the evolution of long-distance migration. In migratory birds, life history theory predicts that the potential survival costs of migrating longer distances should be balanced by benefits to lifetime reproductive success, yet quantification of these reproductive benefits in a controlled manner along a large geographical gradient is challenging. We measured a controlled effect of predation risk along a 3350-kilometer south-north gradient in the Arctic and found that nest predation risk declined more than twofold along the latitudinal gradient. These results provide evidence that birds migrating farther north may acquire reproductive benefits in the form of lower nest predation risk.

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[1] The effects of changing ice and atmospheric conditions on the upwelling of deep nutrient-laden waters and biological productivity in the coastal Beaufort Sea were quantified using a unique combination of in situ and remote-sensing approaches. Repeated instances of ice ablation and upwelling during fall 2007 and summer 2008 multiplied the production of ice algae, phytoplankton, zooplankton and benthos by 2 to 6 fold. Strong wind forcing failed to induce upward shifts in the biological productivity of stratified waters off the shelf.

[1] Global gaseous nitrogen (N 2 ) fixation rates may be underestimated and data is lacking from many regions without conspicuous diazotrophic cyanobacteria, such as cold oceans. We estimated N 2 fixation rates at diverse sites in the Canadian Arctic, including the mouth of the Mackenzie River, the offshore Beaufort Sea, Lancaster Sound, Baffin Bay and a river influenced fjord. We also identified potential diazotrophic communities using a targeted survey of the nifH gene. Nitrogen fixation rates ranged from 0.02 nmol N L À1 d À1in Baffin Bay to 4.45 nmol N L À1 d À1 in the Mackenzie River plume. Sequences recovered from the nifH gene survey belonged mainly to Cluster III, a group of nifH sequences associated with diverse microorganisms, with some a-and g-proteobacteria nifH genes at most sites. Cyanobacteria nifH genes with best matches to Nostocales, which are common in Arctic freshwaters, were recovered from the marine Beaufort Sea. The geographic pattern of N 2 fixation rates and nifH gene identities suggest that the Mackenzie River is the source of a diazotrophic community that contributes new nitrogen to the nitrogen-depleted surface waters of the Beaufort Sea. This first record of N 2 fixation at high latitudes refines our understanding of the global nitrogen budget.