Alexander M. Milner

Island biogeographical theory: can it be used to predict lotic recovery rates?

Classic island biogeographic theory predicts that equilibrium will be reached when immigration and extinction rates are equal. These rates are modified by number of species in source area, number of intermediate islands, distance to recipient island, and size of intermediate islands. This general model has been variously modified and proposed to be a stochastic process with minimal competitive interaction or heavily deterministic. Predictive models of recovery (regardless of the end point chosen) have been based on the appropriateness of the MacArthur-Wilson models.Because disturbance frequency, severity, and intensity vary in their effect on community dynamics, we propose that disturbance levels should first be defined before evaluating the applicability of island biogeographical theory. Thus, we suggest a classification system of four disturbance levels based on recovery patterns by primary and secondary succession and faunal organization by primary (invasion of vacant areas) and secondary (remnant of previous community remains) processes.Level 1A disturbances completely destroy communities with no upstream or downstream sources of colonizers, while some component of near surface interstitial or hyporheic flora and fauna survive level 1B disturbances. Recovery has been reported to take from five years to longer than 25 years, when most invading colonists do not have an aerial form.Level 2 disturbances destroy the communities but leave upstream and downstream colonization sources (level 2A) and, sometimes, a hyporheic pool of colonizers (level 2B). Recovery studies have indicated primary succession and faunal structuring patterns (2A) with recovery times of 90–400 days or secondary succession and faunal structuring patterns (2B) with recovery times of 40–250 days.Level 3 disturbances result in reduction in species abundance and diversity along a stream reach; level 4 disturbances result in reduction of abundance and diversity in discrete patches. Both disturbance types lead to secondary succession and secondary faunal organization. Recovery rates can be quite rapid, varying from less than 10 days to 100 or more days.We suggest that island biogeographical models seem appropriate to recovery by secondary processes after level 3 and 4 disturbances, where competition may be an important organizing factor, while models of numerical abundance and resource tracking are probably of better use where community development is by primary succession (levels 1 and 2).Development of predictive recovery models requires research that addresses a number of fundamental questions. These include the role of hydrologic patterns on colonization dynamics, the role of nonaerial colonizers in recovery from level 1 disturbances, and assessment of the impact of changes in the order of invasion by colonizers of varying energetic efficiencies. Finally, we must be able to assemble these data and determine whether information that guides community organization at one level of disturbance can provide insights into colonization dynamics at other levels.

The effects of climatic fluctuations and extreme events on running water ecosystems

Most research on the effects of environmental change in freshwaters has focused on incremental changes in average conditions, rather than fluctuations or extreme events such as heatwaves, cold snaps, droughts, floods or wildfires, which may have even more profound consequences. Such events are commonly predicted to increase in frequency, intensity and duration with global climate change, with many systems being exposed to conditions with no recent historical precedent. We propose a mechanistic framework for predicting potential impacts of environmental fluctuations on running-water ecosystems by scaling up effects of fluctuations from individuals to entire ecosystems. This framework requires integration of four key components: effects of the environment on individual metabolism, metabolic and biomechanical constraints on fluctuating species interactions, assembly dynamics of local food webs, and mapping the dynamics of the meta-community onto ecosystem function. We illustrate the framework by developing a mathematical model of environmental fluctuations on dynamically assembling food webs. We highlight (currently limited) empirical evidence for emerging insights and theoretical predictions. For example, widely supported predictions about the effects of environmental fluctuations are: high vulnerability of species with high per capita metabolic demands such as large-bodied ones at the top of food webs; simplification of food web network structure and impaired energetic transfer efficiency; and reduced resilience and top-down relative to bottom-up regulation of food web and ecosystem processes. We conclude by identifying key questions and challenges that need to be addressed to develop more accurate and predictive bio-assessments of the effects of fluctuations, and implications of fluctuations for management practices in an increasingly uncertain world.

Glacier shrinkage driving global changes in downstream systems

Glaciers cover ∼10% of the Earth’s land surface, but they are shrinking rapidly across most parts of the world, leading to cascading impacts on downstream systems. Glaciers impart unique footprints on river flow at times when other water sources are low. Changes in river hydrology and morphology caused by climate-induced glacier loss are projected to be the greatest of any hydrological system, with major implications for riverine and near-shore marine environments. Here, we synthesize current evidence of how glacier shrinkage will alter hydrological regimes, sediment transport, and biogeochemical and contaminant fluxes from rivers to oceans. This will profoundly influence the natural environment, including many facets of biodiversity, and the ecosystem services that glacier-fed rivers provide to humans, particularly provision of water for agriculture, hydropower, and consumption. We conclude that human society must plan adaptation and mitigation measures for the full breadth of impacts in all affected regions caused by glacier shrinkage.

Nutrient uptake controls and limitation dynamics in north-east Greenland streams

ABSTRACT Permafrost thaw induced by climate change will cause increased release of nutrients and organic matter from the active layer to Arctic streams and, with increased water temperature, will potentially enhance algal biomass and nutrient uptake. Although essential for accurately predicting the response of Arctic streams to environmental change, knowledge of nutrient release on current Arctic in-stream processing is limited. Addressing this research gap, we quantified nutrient uptake of short-term releases of NO3−, PO43- and NH4+ during peak snowmelt season in five streams of contrasting physiochemical characteristics (from unstable, highly turbid to highly stable, clear-water systems) in north-east Greenland to elucidate the major controls driving nutrient dynamics. Releases were plus or minus acetate to evaluate uptake dynamics with and without a dissolved organic carbon source. To substantiate limiting nutrients to algal biomass, nutrient-diffusing substrates were installed in the five streams for 16 days with NH4+, PO43- or NH4+ + PO43- on organic and inorganic substrates. Observed low uptake rates were due to a combination of low nutrient and DOC concentrations, combined with low water temperature and primary producer biomass, and substantial variation occurred between streams. N was found to be the primary limiting nutrient for biofilm, whilst streams displayed widespread PO43- limitation. This research has important implications for future changes in nutrient processing and export in Arctic streams, which are predicted to include increased nutrient uptake rates due to increased nutrient availability, warmer water temperatures and increased concentration of labile carbon. These changes could have ecosystem and landscape-wide impacts.

Longitudinal distribution of macroinvertebrates in snowmelt streams in northeast Greenland: understanding biophysical controls

In a changing climate, Arctic streams are expected to show more influence from snowmelt, rainfall and groundwater, and less domination from glacial meltwater sources. Snowmelt streams are characteristic features of Arctic ecosystems, yet our current understanding of longitudinal patterns in benthic macroinvertebrate assemblages in these systems is limited when compared to glacier-fed systems. This study characterised longitudinal patterns of macroinvertebrate communities in snowmelt streams in northeast Greenland to provide novel insights into Arctic stream communities as dominant water sources shift with climate change. Benthic macroinvertebrates and environmental variables were sampled at three sites along five streams. Taxa diversity, evenness and abundance were expected to increase with distance from the stream source due to enhanced channel stability and warmer water temperature. This expectation for diversity and evenness was found in two streams, but abundance was up to ten times higher at the upstream sites compared to downstream, where biofilm biomass and ionic load were also highest. Here communities were largely dominated by the genus Eukiefferiella (Chironomidae). In the other three streams, no clear pattern in longitudinal macroinvertebrate community composition was evident due to low channel stability along the entire stream length. This study highlights the considerable variation in macroinvertebrate zonal distribution between snowmelt streams in northeast Greenland. A change towards more snowmelt-dominated streams in the Arctic could lead to shifts in the longitudinal organisation of macroinvertebrate community assemblages and the dominant species as a function of channel stability characteristics.

Controls on Arctic glacier-fed river water temperature

ABSTRACT The impact of climate change on Arctic rivers is expected to be severe. There is therefore a need for greater understanding of Arctic river temperature processes. This study quantifies the spatio-temporal variability of water temperatures in the Kårsa River, Sweden. Water temperature was monitored over two summers within the main proglacial channel and within braids fed by different sources. Longitudinal and lateral temperature patterns were assessed in relation to prevailing hydro-meteorology. Temperature metrics in the main channel increased with distance downstream but were moderated by a large lake, while temperatures in the braids were dependent upon channel source. The high temperature standard deviation and inter-site differences within the braids highlight the importance of braided channels for creating thermal habitat heterogeneity. Temperatures were dependent on hydro-meteorological conditions, with sensitivity to air temperature maximized during cooler, rainy conditions. These results shed new light on Arctic river temperature patterns and their controlling processes.

River ecosystem resilience to extreme flood events

Abstract Floods have a major influence in structuring river ecosystems. Considering projected increases in high‐magnitude rainfall events with climate change, major flooding events are expected to increase in many regions of the world. However, there is uncertainty about the effect of different flooding regimes and the importance of flood timing in structuring riverine habitats and their associated biotic communities. In addition, our understanding of community response is hindered by a lack of long‐term datasets to evaluate river ecosystem resilience to flooding. Here we show that in a river ecosystem studied for 30 years, a major winter flood reset the invertebrate community to a community similar to one that existed 15 years earlier. The community had not recovered to the preflood state when recurrent summer flooding 9 years later reset the ecosystem back to an even earlier community. Total macroinvertebrate density was reduced in the winter flood by an order of magnitude more than the summer flood. Meiofaunal invertebrates were more resilient to the flooding than macroinvertebrates, possibly due to their smaller body size facilitating greater access to in‐stream refugia. Pacific pink salmon escapement was markedly affected by the winter flood when eggs were developing in redds, compared to summer flooding, which occurred before the majority of eggs were laid. Our findings inform a proposed conceptual model of three possible responses to flooding by the invertebrate community in terms of switching to different states and effects on resilience to future flooding events. In a changing climate, understanding these responses is important for river managers to mitigate the biological impacts of extreme flooding effects.

Spatio-temporal dynamics of macroinvertebrate communities in northeast Greenlandic snowmelt streams: Macroinvertebrate community dynamics in Greenlandic snowmelt streams

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