Predictable predator prey scaling - an ecological law?

Some ecologists react with skepticism about the idea of true laws in ecology. So when anything provides a strong and broad relationship between ecological variables, the response is often some combination of surprise, excitement, and disbelief. It’s not unexpected then that a new paper in Science - The predator-prey power law: Biomass scaling across terrestrial and aquatic biomes – has received that reaction, and a fair amount of media coverage too.

Figure 1 from Hatton et al. 2015. "Predators include lion, hyena, and other large carnivores (20 to 140 kg), which compete for large herbivore prey from dik-dik to buffalo (5 to 500 kg). Each point is a protected area, across which the biomass pyramid becomes three times more bottom-heavy at higher biomass. This near ¾ scaling law is found to recur across ecosystems globally."

Ian Hatton and co-authors present robust evidence that across multiple ecosystems, predator biomass scales with prey biomass by a power law with an exponent of ~0.75. This suggests that ecosystems are typically bottom heavy, with decreasing amounts of predator biomass added as more prey biomass is added. The paper represents a huge amount of work (and is surprisingly long as Science papers typically go): the authors compiled a huge database from 2260 communities, representing multiple ecosystems (mammals, plants, protists, ectotherm, and more)(Figure below). Further, the same scaling relationship exists between community biomass and production, suggesting that production drops off as communities increase in density. This pattern appears consistently across each dataset.

Figure 5 from Hatton et al. 2015. "Similar scaling links trophic structure and production.
Each point is an ecosystem at a period in time (n = 2260 total from 1512 locations) along a biomass gradient. (A toP) An exponent k in bold (with 95% CI) is the least squares slope fit to all points n in each row of plots..."

Their analysis is classic macroecology, with all the strengths and weaknesses implicit. The focus is unapologetically on identifying general ecological patterns, with the benefit of large sample sizes, cross system analysis, and multiple or large spatial scales. It surpasses this focus only on patterns by exploring how this pattern might arise from simple predator-prey models. They demonstrate that, broadly, predator biomass can have the same scaling as prey production, which they show follows the 3/4 power law relationship. As for why prey production follows this rule, they acknowledge uncertainty as to the exact explanation, but suggest density dependence may be important.

Their finding is perhaps more remarkable because the scaling exponent has similarities to another possible law, metabolic scaling theory, particularly the ~0.75 exponent (or perhaps ~2/3, depending on who you talk to). It’s a bit difficult for me, particularly as someone biased towards believing in the complexities of nature, to explain how such a pattern could emerge from vastly different systems, different types of predators, and different abiotic conditions. The model they present is greatly 

Peter Yodzis' famous food web for
the Benguela ecosystem.

simplified, and ignores factors often incorporated into these models, such as migration between systems (and connectivity), non-equilibrium (such as disturbance), and prey refuges. There is variation in the scaling exponent, but it is not clear how to evaluate a large vs. small difference (for example, they found (section M1B) that different ways of including data produced variation of +/- 0.1 in the exponent. That sounds high, but it’s hard to evaluate). Trophic webs are typically considered complicated – there are parasites, disease, omnivores, cannibalism, changes between trophic levels with life stage. How do these seemingly relevant details appear to be meaningless?

There are multiple explanations to be explored. First, perhaps these consistent exponents represent a stable arrangement for such complex systems or consistency in patterns of density dependence. Consistent relationships sometimes are concluded to be statistical artefacts rather than actually driven by ecological processes (e.g. Taylor’s Law). Perhaps most interestingly, in such a general pattern, we can consider the values that don’t occur in natural systems. Macroecology is particularly good at highlighting the boundaries on relationships that are observed in natural systems, rather than always identifying predictable relationships. The biggest clues to understanding this pattern maybe in finding when (or if) systems diverge from the 0.75 scaling rule and why.

Ian A. Hatton, Kevin S. McCann, John M. Fryxell, T. Jonathan Davies, Matteo Smerlak, Anthony R. E. Sinclair, Michel Loreau. The predator-prey power law: Biomass scaling across terrestrial and aquatic biomes. Science. Vol. 349 no. 6252. DOI: 10.1126/science.aac6284

Invasion of the Beavers

Guest post by John Cherkas

Fifty years ago, Dr. Walter Howard presented his thoughts on invasive mammals at a symposium on colonizing (invasive) species, which was later turned into the volume "The Genetics of Colonizing Species." He speculated on the nature of predator-prey interactions, population growth limits and habitat disruptions. His ideas still resonate, but how well do they match up with a certain invasive mammal today.

May I bring your attention to some invasive beavers? Our national creature has been making quite a mess is the Southern most reaches of the Western Hemisphere. In the 1940s, Argentina was seeking economic improvements and imported beavers, mink and muskrat to Tierra del Fuego in an attempt to establish a fur trade. That fur trade didn't turn out as expected. Within a few years, beavers had colonized the entire island and were soon crossing channels to reach other Chilean islands, including Cape Horn, a UNESCO Biosphere Reserve.

Angry beaver -roar!

The ecological effects have been pretty well researched recently by Dr. Christopher B. Anderson. In seeing if beavers behave differently in their new habitat than back home, he’s been finding a few differences in the environment and beavers here. One of the most obvious changes is that the beaver colonies are at least twice as dense in Cape Horn. Is this for lack of predators or an abundance of food? So far, I couldn’t say, but I’d lean toward the latter. The Cape Horn forests are entirely southern beeches, which provide ample resources for the beavers’ engineering projects.

But how disruptive have beavers been to the environment: and environment that has no animal that makes such a massive environmental impact as the beaver. Howard suggested that an animal moving into a habitat where its niche doesn’t exist would have wider impacts than one who’s niche does exist. It’s fairly clear that the beaver’s landscaping projects is not something that other animals (except humans) partake in.

In the beaver situation this ecological disruption holds true. The floral assemblage in Cape Horn has never had to deal with beaver-like behaviour. The beavers foraging and building habits prevent forest regrowth, and provide a pathway for other plants to invade. It seems this beaver introduction might be a good example of invasional meltdown. The Chilean archipelago is home to quite a few invasive species already, and this synergistic effect is definitely concerning.

All the beaver-induced worries come with a grave concern for the natural environment. Cape Horn is referred to as pristine quite a bit by Anderson. Is this the place to have a deep political, socio-economic discussion about “pristine” environments? No, not today; you’ll have to read elsewhere for that. Cape Horn is certainly already at risk from invasive species. Beavers have a tremendous impact on the ecological structure of streams and forests. I am certainly one to wonder whether the eradication effort can be truly successful and both removing the beavers and reversing the environmental changes.

I surely hope that the environmental disruption can be reversed. Unfortunately we cannot look back to Howard to speculate on what happens when we remove an alien species. Just fifty years ago, species invasions were seen as a great research opportunity, not something to be extensively managed or eradicated.

Further Reading:

C.B. Anderson et al. (2006), The effects of invasive North American beavers on riparian plant communities in Cape Horn, Chile. Do exotic beavers engineer differently in sub-Antarctic ecosystems? Biological Conservation, 128: 467-474.

doi:10.1016/j.biocon.2005.10.011

C. Choi, (2008) Tierra del Fuego: the beavers must die. Nature 453: 968. doi:10.1038/453968a

Waste not, want not? How human food waste changes ecosystems

Daniel Oro, Meritxell Genovart, Giacomo Tavecchia, Mike S. Fowler and Alejandro Martínez-Abraín. 2013. Ecological and evolutionary implications of food subsidies from humans. Ecology Letters. Volume 16, Issue 12, pp 1501–1514. DOI: 10.1111/ele.12187

Humans have always been connected to their environment, directly and indirectly. Ecologists in particular, and people in general, have been thinking about the causes and consequences of these connections for hundreds of years. One form of interaction results when human food resources become available to other animals – for example, through waste dumps, crop waste, fishery by-catches, bird feeders, or road kill. Starting with middens and waste piles in early human settlements, our food waste has always passed t

Garbage dump in India.

o other species. And while rarely considered compared to human impacts like habitat destruction and climate change, a new review by Daniel Oro and colleagues argues that these subsidies have shaped ecosystems around the globe.

Human food waste (aka subsidies) may come from a variety of human activities, with the three most prominent being crop residuals (remnants of harvest remaining on fields), waste dumps, and fishery discards (by-catch thrown overboard). Each of these forms of subsidy occurs globally and large numbers of species rely partially or completely on them for food. For example, dumps are global in distribution, and contain enough edible waste to attract 20-30% of all mammal and bird in a region (particularly omnivorous and carnivorous species). Crop residues usually attract herbivorous or granivorous species (particularly birds), while fisheries’ waste alter marine ecosystems. Eight percent of all catch (~7 million tonnes!) is simply released back into the ocean, and this supplements species across the food web, including half of all seabirds.

Food waste from human activities may not seem so terrible – after all, they are predictably available, easy to access, fast to forage, and can lead to increased condition and fertility among species that take advantage of them. For example, seabirds foraging among fishing boats for by-catch take advantage of the predictability of boat (and food) appearances and as a result have decreased foraging time and areas, higher individual fitness and reproductive success, and ultimately increased population growth. But the authors suggest that these benefits must be considered in a more complicated web of interactions. After all, human food subsidies tend to be much more predictable than natural sources of food and quickly have large effects spanning from individuals, communities, ecosystems, and evolutionary pressures.

Individuals often, though not always, experience positive effects from subsidies – increased biomass, fertility, and survival, accompanied by changes in dispersal and ranges. If food waste draws in high densities of individuals, it may be associated with greater disease occurrence, or draw in predators attracted to easy pickings. Populations also often respond positively to food subsidies, and become larger and more stable as food waste availability increases. But this boon for one species can cascade through the food web, and have large negative effects in communities and ecosystems. For example, yellow-legged gulls are found around dumps and fishing trawlers, taking advantage of the quantities of food available there, and as a result have increased greatly in population. The downside is that in turn these larger populations increase predation pressure on other vulnerable seabird species. Seabirds in particular can create complicated interactions between human food waste and far-flung ecosystems, connecting as they do both terrestrial and marine systems, moving nutrients, pollution, and calories between systems and through trophic levels.

Snow goose exclosure in northern Canada.
Only the small green rectangle has avoided goose grazing.

A famous example of the unexpected consequences of waste subsidies is the snow goose (Chen chen caerulescens). Snow geese have moved from feeding predominantly on marsh plants to landing en masse in farmers’ fields to feed on grain residues. This new and widespread source of food lead to a population boom, and the high numbers of geese stripped away the vegetation in the arctic habitats where they summer and breed. Agriculture food subsidies in southern habitats were felt far away in the arctic, as migratory snow geese tied these systems and food webs together. Though snow geese are unlikely to lose their new source of food, other animals may face plummeting populations or extinctions if food sources disappear. Until the 1970s, in Yellowstone, grizzly bears fed nearly exclusively at a local dump that then closed: the result was both increased mortality and rapid increases in foraging distances and behaviours.

Finally, and of most concern, food waste subsidies can alter the selective pressures a population faces. Species that become reliant on dumps or fields for food may experience changes in selective pressures, leading to selection for traits necessary to exploit these subsidies, and a loss of genotypic/phenotypic variation from the population. Changes in selective pressure change with the situation, of course. In the case of Yellowstone, the dump closure and loss of food source seemed to have large effects on traits important for sexual attractiveness in males, suggesting potential effects on reproductive success. In the best known (and my favourite) example of the selective effect of human food waste, dogs eventually were domesticated from wolf ancestors. Of course dumps can also relax selective pressures, if they allow individuals in poor condition (juveniles, the elderly) to successfully feed and reproduce.

Though food waste subsidies are clearly important and can have wide ranging effects, it is worth noting that the effect and importance of food subsidies is context-dependent. Studies seem to indicate that effects are greatest when food is low naturally or habitat quality is poor; in high quality systems, food waste may only be used by juveniles or individuals in poor condition. Unfortunately, as humans degrade natural habitats, subsidies are only likely to increase in importance as a food source for species. The extent and effects of human food waste are yet another legacy of the global alterations the human species has made. Unfortunately, like so many of the changes we have made, the issues are complex and transcend political and regional boundaries. Practices in one system or nation are tied to effects in another nation, and this complexity can make it difficult to monitor and measure the effects of subsidies as thoroughly as is necessary. This review from Oro et al. certainly makes a case for why our garbage needs to receive more attention.

One example of the ecosystem wide effects of subsidies: here, fisheries inputs.

Can intraspecific differences lead to ecosystem differences?

Sara Lindsay Jackrel and J. Timothy Wootton. 2013. Local adaptation of stream communities to intraspecific variation in a terrestrial ecosystem subsidy. Ecology. Online early.

It’s funny how complex outcomes can arise from simple realizations. For example, it is plausible that when there are differences among individuals of a species (like when local populations are adapted to the local environment), these could implications for function on the ecosystem scale. But while there is increasing evidence for the importance of intraspecific variation for ecological interactions within communities, the question of how intraspecific diversity scales up to ecosystem functioning is still ambiguous.

Sara Jackrel and Timothy Wootton explore this question in “Local adaptation of stream communities to intraspecific variation in a terrestrial ecosystem subsidy”. The basis for their study was simple: local adaptation is common, and populations/genotypes/ecotypes tend to be best adapted to the particular conditions of their locale. For example, “spatial variation in prey and predators can lead to a geographic mosaic of co-evolutionary interactions”. Further, these localized interactions can affect the greater ecosystem, if individuals or materials move between ecosystem boundaries.

In particular, the authors note that there is evidence that tree species composition riverside can alter the composition of the local aquatic community. This occurs via leaf litter fluxes into the river: the type and amount of leaf litter that falls into streams varies, and so the type of macroinvertebrates in the recipient stream also varies in response. These macroinvertebrates break down the leaf litter via shredding, collecting, and filtering, playing an important role in nutrient cycles. Leaf litter is carried from a given tree by wind or water and may decompose near or far away, creating a connection between ecosystems. The question then is whether macroinvertebrate compositional shifts will occur in response to intraspecific differences in leaf (i.e. trees), and what the implications might be for ecosystem functions such as leaf decomposition. To explore this question, Jackrel and Wootton performed reciprocal transplants of leaf litter material between eight sites along rivers in the Olympic Peninsula of Washington.

All eight of these sites were early successional forests dominated by red alder. The authors collected fresh leaves from alder trees, bagging leaves from each tree separately. These bags of leaves were either placed in the river adjacent to the trees they were taken from, or in a more distant site. Non-adjacent sites were either in the same river as the home site, or in different river all together. Leaf packs were weighed before and after spending 17-18 days in the river. This would allow comparison of how decomposition rates vary between home and away sites, and between home and away rivers.

Their results suggested a few interesting things. First, the identity of a tree affects the rate of decomposition of its leaves: individual alder trees’ leaves were highly variable in the rate of decomposition. Second, the combined identities of trees at a site seem to have affected the composition of the decomposer community at the home river site: put leaves from that site in another river with a new community of decomposers, and the decomposition rate drops significantly. In general, leaves decomposed significantly more rapidly when in their home river, regardless of whether at the home site or elsewhere along the river. But if they put leaves upstream from the home site, but in the same river, the rate of decomposition also dropped. Upstream decomposer communities were apparently much worse at breaking down leaves from novel communities of alders. However, if you put the leaves in sites downstream from home, the decomposition rates are not significantly different than in the home site. This is likely because of the directional movement of a river, such that downstream locations receive leaf litter from all upstream sites, and so downstream decomposer communities experience a greater variety of leaf litter than upstream sites. This might lead to upstream sites being more closely adapted to the individual trees in their neighbourhood than downstream sites, which receive inputs from a wide variety of trees. These results suggest that individual differences in trees at different spatial locations can matter, both locally, across trophic levels, and even across ecosystems.

Admittedly there is not a lot you can infer about the mechanisms at play from this preliminary experiment. One interesting follow up would be to measure compositional differences in aquatic macroinvertebrates at very fine scales in correspondence with differences in trees. Another important question is whether these communities differ via phenotypic plasticity, adaptation to local sites, or species sorting. But this paper does hint at one way in differences among individuals can shape local ecosystems and even structure distant ecosystems (e.g. downstream decomposer communities) through fluxes across boundaries. And that is a rather complicated implication from a logical and simple starting point.

Studying Frankenstein: what can we learn from novel ecosystems?

There's been some discussion going around ecolog about an article telling the ecological story of Ascension Island. I should note that the original article is not a great example of science writing; it tries to create conflict that doesn’t exist and lacks a reasonable understanding of ecological theory. There are a couple linked chapters/publications about Ascension Island that make better additions to the story though (1, 2).

Ascension Island is one of those tiny islands first visited by Europeans in the 1600s. Like many young, small, isolated islands (1200 mi to the next nearest island), it was highly depauperate (~25-30 species of plants). Like many such islands, once humans became regular visitors, new species began to make their to way Ascension. The Brits and their love of cultivating and homogenizing particularly altered the island, and they systematically introduced species calculated to provide ecosystem services, aesthetic value, and food.

As a result, Ascension Island changed strikingly – once an island with lowland deserts and a rocky, barren mountainside, the mountain is today known as Green Mountain. The originally depauperate mountain is now lush with three different vegetation zones, a large variety of plants including “banana, ginger, juniper, raspberry, coffee, ferns, fig trees, Cape Yews, and Norfolk Island pines”, and a complex cloud forest. The original article presents this as some inexplicable outcome, but frankly it seems in keeping with existing ecological ideas. Under island biogeography, if you decrease the distance from an island to the mainland (including via human-aided dispersal), diversity should increase. Given the massive number of species that were introduced, and the coddling they received to aid their establishment, heightened diversity is hardly a surprise. And though the original article suggests that shared evolutionary history is necessary for complex ecosystems, coevolution is hardly a requirement for a functioning ecosystem to develop. Species may be able to coexist despite lacking a shared history--niches may not be filled as tightly as in a long-established, coevolved community, but invasive species research in general should have taught us that novel species combinations can easily occur. Secondly, many of the introduced species on the island are from the same part of the world and likely do share evolutionary history.

The mountain before and after. From Catling & Stroud.

from Hobbs et al. 2006

I hadn't given much thought before to the concept of “novel ecosystems” and it has received little attention from the ecological literature (excepting the odd papers, and much more attention from a conservation and management angle). Ascension is a particularly striking example of how human modification leads to ecosystems which are entirely different from anything that has ever been present on the planet. Novel ecosystems have been defined in a number of ways. Generally, they are synthetic ecosystems that include conditions and combinations of organisms never before in existence, and do not depend on human maintenance to persist (as agriculture fields would). Novel ecosystems may be considered to be the outcome of abandonment of human managed systems or else the degradation of existing systems through human activities and invasion (figure). Of course there are incredibly few ecosystems that aren’t affected in some way by human activities (especially in this age of intentional and unintentional human-mediated species introductions), but it is the truly unique ones that are particularly interesting.

There are at least two ways to approach novel ecosystems. One approach is parallel with invasive species and conservation research, and in fact these research areas overlap a fair amount. This is the way in which most research on novel ecosystems seems to be framed. Novel ecosystems carry many of the same issues about making value judgments as invasive species research, and issues of management and whether novel ecosystems can or should be returned to their original state dominate. For example, the conflict between maintaining alpha (island) and gamma (global) diversity exists on Ascension Island– modern, invaded Ascension Island provides greater diversity and ecosystem functioning (erosion control, food, temperature moderation, habitat) than the original barren landscape. But the original endemic species, not surprisingly, have gone extinct or are increasingly at risk.

But focusing solely on these difficult value-laden questions seems to have been at the cost of exploring the value of novel ecosystems as a study system. The most interesting examples of novel ecosystems are not simply modified or invaded ecosystems, but ecosystems that truly never existed before. Like post-shale dump landscapes in Scotland, where the refuse from mining is now host to unique grasslands that act as refugia for locally rare species; or the San Francisco Bay, which now is utterly unrecognizable compared to historical descriptions due to heavy invasion; or urban ecosystems with their unique habitats and issues; or even the habitat and connectivity created by stone fences which now occur on most continents. The questions here aren't always about invasion and management, but instead focus on what the new community looks like. How do novel communities assemble, what processes dominate (mass effects, environmental filtering, competition, predation, etc, etc)? How does ecosystem function relate to the community that assembles? Most BEF research after all, is focused on more traditional ecosystems. What leads to stability in a novel ecosystem, or are they stable at all? They can function is an example of highly unfortunate but also highly informative ‘natural’ experiments for ecologists. But at the moment, if you search for "novel ecosystems" on Google Scholar, the title words are "management", "conservation", "restoration" or "invasion". Actually, there probably are ecologists doing work on novel ecosystems from a purely ecological perspective, but this work gets grouped with  disturbance, invasion, and urban ecology: it just remains to consider them in a more unified fashion. If the conversation remains focused only on the conservation issues (as the discussion on ecolog seemed to shift to rapidly), it just seems like we're limiting ourselves a little.

Grassland diversity increases stability across multiple functions

As ecological systems are altered with cascading changes in diversity, the oft-asked question is: does diversity matter for ecosystem function? This question has been tested a multitude of times, with the results often supporting the idea that more diverse assemblages provide greater functioning (such as productivity, nutrient cycling, supporting greater pollinator abundance, etc.). Besides greater functioning, scientists have hypothesized that more diverse systems are inherently more stable. That is, the functions communities provide remain more constant over time compared with less diverse systems, which may be less reliable.

While the relationship between diversity and stability has been tested for some functions, Proulx and colleagues examined the stability of 42 variables over 7 years across 82 experimental plots planted with either 1, 2, 4, 8, 16 or 60 plant species in Jena, Germany. They examined patterns of variation (and covariation) in the functions and found that many show lower variation over time in plots with more plant species. Greater stability was found at many different trophic levels including plant biomass production, the abundance and diversity of invertebrates and the abundance of parasitic wasps -which indicate more complex food webs. They also found greater stability in gas flux, such as carbon dioxide. Despite the greater stability in these measures of above-ground functions, below ground processes, such as earthworm abundance and soil nutrients, were not less variable in high diversity plots.

How ecosystems function is of great concern; these results show that more diverse plant communities function more stably and reliably than less diverse ones. The next step for this type of research should be to address what kind of diversity matters. A greater number of species means more different kinds of species, with differing traits and functions. What aspect of such functional differences determine stability of ecosystem function?

This is an exciting paper that continues to highlight the need to understand how community diversity drives ecosystem function.

Proulx, R., Wirth, C., Voigt, W., Weigelt, A., Roscher, C., Attinger, S., Baade, J., Barnard, R., Buchmann, N., Buscot, F., Eisenhauer, N., Fischer, M., Gleixner, G., Halle, S., Hildebrandt, A., Kowalski, E., Kuu, A., Lange, M., Milcu, A., Niklaus, P., Oelmann, Y., Rosenkranz, S., Sabais, A., Scherber, C., Scherer-Lorenzen, M., Scheu, S., Schulze, E., Schumacher, J., Schwichtenberg, G., Soussana, J., Temperton, V., Weisser, W., Wilcke, W., & Schmid, B. (2010). Diversity Promotes Temporal Stability across Levels of Ecosystem Organization in Experimental Grasslands PLoS ONE, 5 (10) DOI: 10.1371/journal.pone.0013382

Enhanced biodiversity-ecosystem function relationships in polluted systems

*note: this text was adapted from an Editor's Choice I wrote for the Journal of Applied Ecology.

In this era of species loss and habitat degradation, understanding the link between biodiversity and functioning of species assemblages is a critically important area of research. Two decades of research has shown that communities with more species or functional types results in higher levels of ecosystem functioning, such as nutrient processing rates, carbon sequestration and productivity, among others. This research has typically used controlled experiments that standardize environmental influences and manipulate species diversity. However, a number of people have hypothesized that biodiversity may be even more important for the maintenance of ecosystem functioning during times of environmental stress or change rather than under stable, controlled conditions. It is during these times of environmental change that preserving ecological function is most important, as changes in function can have cascading effects on other trophic levels, compounding environmental stress. Therefore, explicitly testing how biodiversity affects function under environmental stress can help to inform management decisions.

Image from Wikimedia commons

In a recent paper in the Journal of Applied Ecology, Li and colleagues examine how algal biodiversity influences productivity in microcosms with differing cadmium concentrations. Cadmium (Cd) is a heavy metal used in a number of products and industrial processes, but it is toxic and Cd pollution is a concern for human populations and biological systems, especially aquatic communities. This is especially true in nations currently undergoing massive industrial expansion. In response to concerns about Cd pollution effects on aquatic productivity, Li et al. used algal assemblages from single species monocultures to eight species polycultures grown under a Cd-free control and two concentrations of Cd, and measured algal biomass.

Their results revealed that there was only a weak biodiversity-biomass relationship in the Cd-free teatment, which the authors ascribed to negative interactions offsetting positive niche partitioning. In particular, those species that were most productive in their monocultures were the most suppressed in polycultures. However, in microcosms with Cd present there were positive relationships between diversity and biomass. They attribute this to a reduction in the strength of competitive interactions and the opportunity for highly productive species to persist in the communities.

While a plethora of experiments generally find increased ecosystem function with greater diversity, Li et al.’s research indicates that the effect of biodiversity on function may be even more important in polluted systems. If this result can be duplicated in other systems, then this gives added pressure for management strategies to maintain maximal diversity as insurance against an uncertain future.

Li, J., Duan, H., Li, S., Kuang, J., Zeng, Y., & Shu, W. (2010). Cadmium pollution triggers a positive biodiversity-productivity relationship: evidence from a laboratory microcosm experiment Journal of Applied Ecology, 47 (4), 890-898 DOI: 10.1111/j.1365-2664.2010.01818.x

To intervene or not to intervene: this is a real question

Should land managers actively manipulate the structure and function of ecosystems within protected areas? Is intervention appropriate to protect or maintain native biodiversity and natural processes in areas such as national parks and wilderness areas? These are the questions that stem from a new paper by Richard Hobbs and others in Frontiers in Ecology and the Environment. US national parks and wilderness areas have legislative mandates to maintain ‘naturalness’, but what does this mean in the context of dynamic ecosystems with current and future changes including invasions by nonnative organisms and climate change?

Hobbs and his colleagues challenge concepts of naturalness and propose several ‘guiding principles’ for stewards of national parks and wilderness. They suggest that more useful concepts for managing protected areas relate to ecological integrity and resilience. Concepts of ecological integrity have been adopted by Parks Canada and relate to maintaining ecosystem components. Resilience concepts focus on the ability of a system to “absorb change and persist” without undergoing a “fundamental loss of character”. While maintaining ecological integrity in the face of global changes may - by definition - require protection of species, maintaining ecological resilience tends to focus more attention on ecosystem function “over preserving specific species in situ”.

Rather than protecting an area to maintain naturalness, focusing on ecological integrity and resilience acknowledges that a diversity of approaches - from non-intervention to actively managing systems - may be required. The flexibility in this view, demands that conservation planning span gradients of land uses across landscapes. Management objectives and success need to be re-evaluated in an adaptive and experimental framework, which requires careful and robust monitoring.

At The Wilderness Society and specifically here in Montana, these very questions are being wrestled with in terms of forest restoration, fire management, and climate change. Current forest conditions have been shaped by historic logging practices and fire suppression leading to altered structure and function – including increasing the severity of fires. Through active management, including removing small diameter trees and lighting prescribed fires, managers hope to restore forests and fire intensities to conditions more closely resembling those that historically occurred. Much of the research on restoration was conducted in dry forests in the American Southwest where low-severity fires occurred across large areas. However, in the Northern Rockies, many forests were shaped by a ‘mixed severity’ fire regime, where fires crept along the forest floor in some areas and torched trees in others. In many cases, these forests have not been fundamentally altered and need only the return of fire to restore their resilience. In other cases, forests are recovering from past logging practices and may benefit from thinning to restore a fire-resilient structure.

To return to the paper at hand: what is the appropriate level of intervention to maintain ecological integrity and resilience given past forest management and future climate change? If the current forest lacks integrity (novel stand structure) and resilience under a predicted climate of warmer, drier conditions, what is the appropriate level of management? While The Wilderness Society continues to work with diverse partners to answer these questions, one thing is clear: whatever actions take place, they need to be conducted with humility in an experimental framework that includes sufficient ecological monitoring. For the ‘experiment’ to be most helpful, we should maintain adequate hands-off “controls” along with the “treatments” to allow us to gauge the effects of intervention.

Richard J Hobbs, David N Cole, Laurie Yung, Erika S Zavaleta, Gregory H Aplet, F Stuart Chapin III, Peter B Landres, David J Parsons, Nathan L Stephenson, Peter S White, David M Graber, Eric S Higgs, Constance I Millar, John M Randall, Kathy A Tonnessen, Stephen Woodley (2009) Guiding concepts for park and wilderness stewardship in an era of global environmental change. Frontiers in Ecology and the Environment e-View.
doi: 10.1890/090089
http://www.esajournals.org/doi/abs/10.1890/090089