Competitive coexistence, it's all about individuals.

Understanding how species coexist has been the raison d'etre for many ecologists over the past 100 years. The quest to understand and explain why so many species coexist together has really been a journey of shifting narratives. The major road stops on this journey have included searching for niche differences among species -from single resources to multidimensional niches, elevating the role for non-equilibrial dynamics -namely disturbances, and assessing the possibility that species actually differ little and diversity patterns follow neutral process. Along this entire journey, researchers (especially theoreticians) have reminded the larger community that that coexistence is a product of the balance between interactions among species (interspecific) and interactions among individuals within species (intraspecific). Despite this occasional reminder, ecologists have largely searched for mechanisms dictating the strength of interspecific interactions.

Image used under Flickr creative commons license, taken by Tinken

In order for two species to coexist, intraspecific competition must be stronger than interspecific -so sayeth classic models of competition. While people have consistently looked for niche differences that reduce interspecific competition, no one has really assessed the strength of intraspecific competition. Until now that is. In a recent paper in Science, Jim Clark examines intra- vs interspecific interactions from data following individual tree performances, across multiple species, for up to 18 years. This data set included annual growth and reproduction, resulting in 226,000 observations across 22,000 trees in 33 species!

His question was actually quite simple -what is the strength of intraspecific interactions relative to interspecific ones? There are two alternatives. First, that intraspecific competition is higher, meaning that among species differences only need to be small for coexistence to occur; or secondly, that intraspecific competition is lower, requiring greater species niche differences for coexistence. To answer this he looked at correlations in growth and fecundity between individuals either belonging to the same or different species, living in proximity to one another. He took a strong positive correlation as evidence for strong competition and a negative or weak correlation as evidence for resource or temporal niche partitioning. What he found was that individuals within species were much more likely to show correlated responses to fluctuating environments, than individuals among species.

This paper represents persuasive evidence that within-species competition is generally extremely high, meaning that to satisfy the inequality leading to coexistence: intra > inter, subtle niche differences can be sufficient. These findings should spur a new era of theoretical predictions and empirical tests as our collective journey to understanding coexistence continues.

Clark, J. (2010). Individuals and the Variation Needed for High Species Diversity in Forest Trees Science, 327 (5969), 1129-1132 DOI: 10.1126/science.1183506

Check out the carnival of evolution and be sure to vote for your favorite blogs

Be sure to check out this month's Carnival of Evolution (number 21) posted at Mauka to Makai. The Carnival is a monthly digest of recent evolutionary musings from around the blogosphere. This month's edition includes a number of interesting posts, as well as one of our posts on what evolution offers conservation.

Also, Research Blogging has announced finalists for various blogs awards. If you are eligible, please vote, there are a lot of great blogs vying for these awards. Also, The EEB and Flow is among the finalists for best biology blog. And to the people we nominated us, thanks again for nominating our blog.

New Tool Reveals Where Ticks Eat Breakfast

You have a much greater chance of getting sick from a tick bite today than you did 30 years ago. But a new tool might allow researchers to better understand why more ticks are making people sick.

“If you’re a health inspector and a bunch of people get food poisoning, the first thing you’d want to know is where they ate last. If you’re a disease ecologist and a bunch of ticks have a pathogen, the first thing you’d want to know is where the ticks ate last,” said Brian Allan, a post-doctoral researcher at the Tyson Research Station in St. Louis.

Allan led a team of researchers in developing a novel technology that probes the genetic contents of ticks’ gut. The tool can determine which wildlife species provided the tick’s last meal and which pathogens came along with that meal.

In the first study to use the new technology, Allan and his colleagues focused on several rapidly emerging diseases transmitted by the lone star tick. These include two pathogens responsible for a potentially fatal bacterial infection known as ehrlichiosis [ur-lick-ee-oh-sis]. In Missouri, over 200 cases of ehrlichiosis were documented last year.

Allan et al.'s study showed that about 80 percent of pathogen-positive ticks had fed on white-tailed deer. They also found that squirrels and rabbits were capable of infecting ticks at a higher rate than deer. However, since the lone star tick feeds on squirrels and rabbits less frequently, they account for a smaller percentage of infection.

Allan and his colleagues hope that the technique will shed light on theoretical questions in the field of ecology. They are especially interested in testing whether biodiversity is good for your health, a hypothesis known as “the dilution effect.”

Allan, B. F., L. S. Goessling, G. A. Storch, and R. E. Thach. 2010. Blood meal analysis to identify reservoir hosts for Amblyomma americanum ticks. Emerging Infectious Diseases 16: 433-440. DOI: 10.3201/eid1603.090911

How can evolution inform conservation decisions?

First of all, let me apologize for the lack of blog posts over the past 2 weeks, I've been busy visiting the Olympics and reading a couple of hundred blogs, judging them for the Research Blogging awards.

The conservation of biological diversity is a major imperative for biologists. International agreements such as the Convention on Biological Diversity and intergovernmental exercises, such as the Millennium Ecosystem Assessment, call upon scientists to provide evidence on the current state of biological diversity and to evaluate solutions for reducing diversity and ecosystem function loss. Critical to these efforts have been the work of ecologists, conservation biologists and ecological economists. However, seemingly missing from the conversation about the state of biodiversity knowledge has been evolutionary biologists. Are they primarily concerned with describing historical processes and mechanisms of biological change, or do they have substantive knowledge and ideas that should be viewed as a critical component of any scheme to conserve biological diversity?

In a recent paper in Evolution, Hendry and a number of coauthors convincingly make the case that evolutionary biology is a necessary component for conservation. Evolution offer four key insights that should inform conservation and policy decisions. First, they point out that evolutionary biologists are in the business of discovering and documenting biodiversity. They are the primary drivers behind long-term, sustained biological collections, because they need to know what exists in order to better understand evolutionary history. With millions of species awaiting scientific discovery, their efforts are critical to measuring biodiversity. But not only are they discovering new species and enumerating them, they are uncovering their evolutionary relationships, which gives conservationists better information about which species to prioritize. What Vane-Wright famously called 'the agony of choice', with limited resources, we need to prioritize some species over others, and their evolutionary uniqueness ought to be a factor. More than this, evolutionary biologists have developed pragmatic tools for inventorying and sharing data on biodiversity at all levels, from genes to species, which is available for prioritization.

The second key insight is that by understanding the causes of diversification, we can better understand and predict diversity responses to environmental and climatic change. By understanding how key functional traits evolve, we can develop predictions about which species or groups of species can tolerate certain perturbations. Further, research into how and why certain evolutionary groups faced extinction can help us respond to the current extinction crisis. For example, the evolutionary correspondence between coevolved mutualists, such as plants and pollinators, can be used to assess the potential for cascading extinctions. These types of analyses can help identify those groups of related species, or those possessing some trait, which make species more susceptible to extinction.

Thirdly, evolution allows for an understanding of the potential responses to human disturbance. Evolutionary change is a critical part of ecological dynamics, and as environment change can result in reduced fitness, smaller population sizes and extinction, evolution offers an adaptive response to these negative impacts. Knowing when and how populations can evolve is crucial. Evolutionary change is a product of genetic variation, immigration, population size and stochasticity, and if the ability to evolve to environmental change is key for persistence, then these evolutionary processes are also key.

Finally, evolutionary patterns and processes have important implications for ecosystem services and economic and human well-being. Both genetic and evolutionary diversity of plant communities has been shown to affect arthropod diversity, primary productivity (including work by me) and nutrient dynamics. Thus understanding how changes in diversity affect ecosystem processes should consider evolutionary processes. Further, exotic species are often cited as one of the major threats to biodiversity, and evolutionary change in exotics has been shown to increase exotic impacts on native species.

All together, these key reasons why evolution matters for conservation, mean that developing sound management plans requires considering evolution patterns and processes. We can use evolution to our benefit only if we understand how evolution shapes current dynamics. The challenge to evolutionary biologists is the same as it was for ecologists perhaps 15 to 20 years ago, to present their understanding and conservation ideas to a broader audience and to engage policy makers. To this end, the authors highlight some recent advances in incorporating evolutionary views into existing biodiversity and conservation programmes –most notably into DIVERSITAS.

Just like ecological processes and dynamics cannot be fully understood without appreciating evolution ancestry or dynamics, developing an extensive, expansive conservation strategies must take into account evolution. I hope that this paper signals a new era of a synthesis between ecology and evolution, which produces precise, viable conservation strategies.

Hendry, A., Lohmann, L., Conti, E., Cracraft, J., Crandall, K., Faith, D., Häuser, C., Joly, C., Kogure, K., Larigauderie, A., Magallón, S., Moritz, C., Tillier, S., Zardoya, R., Prieur-Richard, A., Walther, B., Yahara, T., & Donoghue, M. (2010). EVOLUTIONARY BIOLOGY IN BIODIVERSITY SCIENCE, CONSERVATION, AND POLICY: A CALL TO ACTION Evolution DOI: 10.1111/j.1558-5646.2010.00947.x

Research blogging awards; and thanks

Hi all, nominations for Research Blogging's annual awards will close Feb. 11, so be sure to nominate any research blogs you think deserve consideration. The top prize is $1000, and there are several smaller, field-specific awards as well. A panel of judges (with me being a member) will create a short list of blogs for each category and registered users on Research Blogging will be able to vote for the winners.

And a thank you to whoever nominated the EEB and Flow.

Predator-human conflict: the emergence of a primordial fear?

There is something terrifying and at the same time captivating about the idea of a large, wild, mysterious predator. The very idea that a large predator is near by makes us feel vulnerable. Every year, news stories about wild animal attacks appear in numerous publications and on many television shows. Human death at the fangs or claws of a wild beast is at the heart of many legendary stories and probably sown into the fabric of our being by millennia of ever present risk from large predators. This characteristic of our human experience, I think, dictates our response to animal attacks. Stories of animal attacks are usually concluded with statements about having or attempting to track down and destroy the guilty animal.

Such is the case for three recent animal attacks in Canada. In late October, 2009 in Nova Scotia, a raising 19-year old folk singer was killed by a couple of coyotes while hiking. It is difficult to find meaning in such a horrendous death, but the narrative, told by reporters, was essentially to rest assured that one of the coyotes had been killed and the other was being tracked and would be destroyed. There were two cougar attacks in early January, 2010 in British Columbia, that basically ended with the same reassurance. In the first, a boy was attacked and his pet golden retriever courageously saved his life. A police officer arrived a shot the cougar which was mauling the dog -an obviously legitimate response, and the news story again reassures us that the animal was destroyed. And don't worry the hero dog survived. In the second cougar attack, another boy was attacked, and this time his mother saved his life. But again the story narrative ended by reassuring us that the guilty cougar, and another cat for good measure, were destroyed the next day.

After reading these stories, I asked myself two things. Why is our response to destroy predators that attack? And why do we need to be reassured that this has happened? In defence of the predators, they are just doing what their instincts tell them to do, and most often their only mistake is that they selected their prey poorly. But the reality is that there are only 2-4 cougar attacks per year and only 18 fatalities over the past 100 years. Why do we fear such a low probability event? In contrast, automobile accidents are the leading cause of death in children under 12 in North America. Thousands of people die, and millions injured in car accidents every year in North America. Recently, in Toronto, were I live, 10 pedestrians were killed in 10 days, yet my heart doesn't race when I cross a street. If our fears and responses to human injury and death reflected the actual major risks, we would invoke restrictive rules regarding automobile use.

We believe that we can live with nature in our backyard. But when that close contact results in an animal attack, human fear seems to dictate an irrational response. Do we really expect predators to obey our rules? Can we punish them enough to effectively tame them? We cannot, and I hope that our approaches to dealing with human-animal conflict can better deal with animal attacks, in a way that does not subjugate large predators to whims of our fears.

The evolution of a symbiont

The evolution of negative interactions seems like a logical consequence of natural selection. Organisms compete for resources or view one another as a resource, thus finding ways to more efficiently find and consume prey. However, to me, the natural selection of symbiotic or mutualistic interactions has never seemed as straight forward (expect maybe the case where one species provides protection for the other, such as in ant-plant mutualisms). A specific example is the rise of nitrogen-fixing plants, who supply nutrients to bacteria called rhizobia capable of converting atmospheric nitrogen into forms, such as ammonia, usable to the plant host. Not only has this symbiosis evolved, but has seemed to evolve in very evolutionarily distinct lineages. The question is, what are the mechanisms allowing for this?

In a recent paper, Marchetti and colleagues answer part of the question. They experimentally manipulate a pathogenic bacteria and observe it turning into a symbiont. They transferred a plasmid from the symbiotic nitrogen fixing Cupriavidus taiwanensis into Ralstonia solanacearum and infected Mimosa roots with it. Plasmid transfer among distinct bacteria species is common and referred to horizontal genetic transfer (as opposed to vertical, which is the transfer to daughter cells). The presence of the plasmid caused R. solanacearum to quickly evolve into a root-nodulating symbiont. Two regulatory genes lost function, and this caused R. solanacearum to form nodules and to impregnate Mimosa root cells.

This extremely novel experiment reveals how horizontal gene transfer can supply the impetus for rapid evolution from being a pathogen to a symbiont. More importantly it reveals that sometimes just a few steps are required for this transition and how distantly-related bacterial species can acquire symbiotic behaviors.

Marchetti, M., Capela, D., Glew, M., Cruveiller, S., Chane-Woon-Ming, B., Gris, C., Timmers, T., Poinsot, V., Gilbert, L., Heeb, P., Médigue, C., Batut, J., & Masson-Boivin, C. (2010). Experimental Evolution of a Plant Pathogen into a Legume Symbiont PLoS Biology, 8 (1) DOI: 10.1371/journal.pbio.1000280

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

Timing is everything: global warming and the timing of species interactions

While an obvious affect of climate change will be changes in the distributions or range sizes of species, more insidious and likely more consequential will be how species interactions are affected by changes in the timing of growth and reproduction. These changes in an organism's life cycle, or phenology, can create mismatches between an organism's need and resource availability or the readiness of coevolved partners -such as plants and pollinators.

In an 'Idea and Perspective' paper in Ecology Letters, Louie Yang and Volker Rudolf set out a new framework to examine the effects of phenological shifts on species interactions. They argue that one cannot understand or predict the fitness consequences of a phenology shift without knowing how interacting species' phenologies are also influenced by environmental changes. The consequences of phenological shifts are changes in fitness, and the question is: how would one go about assessing the fitness effects of phenological changes on interactions? This is where this paper really hits its stride. Yang and Rudolf set out a new conceptual framework for studying the fitness consequences of phenological shifts. They make the case that an experimental approach is required to test the three likely scenarios. The first is that there are no changes in phenology -that is, measuring the fitness levels of the two interacting species under stable conditions. Second, you induce an experimental shift in the timing of one of the species. For example, in a plant-herbivore interaction, germinate the plant earlier and when the herbivore normally has access to the plant, the plant will be older. What are the fitness changes associated with this shift? Finally, you can shift the timing of the other species relative to the first. In our example, the herbivore has access to younger plants and again are there fitness consequences?

Yang and Rudolf call the full combination of possible fitness effects, across a number of timing mismatches, 'the ontogeny-phenology landscape'. By mapping fitness changes across this ontogeny-phenology landscape, researchers can offer better predictions, on top of just changes in range size or habitat use, about the possible affects of climate change. The obvious question, and Yang and Rudolf acknowledge this, is how to extend two-species ontogeny-phenology to multi-species communities. Of course, extending two-species interactions to communities is a question that plagues most of community ecology, but I think the solution is that researchers who know their systems often have intuition about the major players, and thus those species where phenology shifts should have disproportionate effects on other species. Such species could be the place to start. Another strategy would be a food web type approach, where species are lumped into broader trophic groups and we ask how shifts in certain trophic groups affect other groups.

Regardless of how to extend this framework to multispecies assemblages, I see this paper as likely to be very influential. It gives researches a new focus and framework, where specific predictions about climate change can be made.

Yang, L., & Rudolf, V. (2010). Phenology, ontogeny and the effects of climate change on the timing of species interactions Ecology Letters, 13 (1), 1-10 DOI: 10.1111/j.1461-0248.2009.01402.x

Plant genotypic diversity supports pollinator diversity

Research over the past 20 years has shown that plant communities with greater diversity maintain higher productivity, greater stability and support more diverse arthropod assemblages. More recently, several experiments have shown that interspecific diversity (namely genotypic differences) also affects community functioning. Pollination is often considered an essential function, and does plant genotypic diversity affect pollinator diversity and frequency?

In a recent paper in PLoS ONE, Genung and colleagues test whether plant genotypic diversity affects pollinator visits. They use an experimental system set-up by Greg Crutsinger that combines multiple genotypes of the goldenrod, Solidago altissima, and record pollinator visits over two years. Experimental plots contained 1, 3, 6, or 12 genotypes of S. altissima. After accounting for differences in abundance, Genung et al. show that as genotypic diversity increases, both pollinator richness and number of visits to the plot significantly increase. This increase is greater than expectations of randomly simulated assemblages combining proportional pollinator visits from monocultures.

The previous research at the species-level has made a persuasive rationale to protect species diversity in order to maintain ecosystem functioning. Now, research like this is making a case that there are consequences for not explicitly considering genetic diversity in conservation planning and habitat restoration.

Genung, M., Lessard, J., Brown, C., Bunn, W., Cregger, M., Reynolds, W., Felker-Quinn, E., Stevenson, M., Hartley, A., Crutsinger, G., Schweitzer, J., & Bailey, J. (2010). Non-Additive Effects of Genotypic Diversity Increase Floral Abundance and Abundance of Floral Visitors PLoS ONE, 5 (1) DOI: 10.1371/journal.pone.0008711