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

Organic farming and natural enemy evenness

The basic reality of agricultural activity is that it reduces biological diversity, and these reductions in diversity potentially impact ecosystem services. But do some agricultural practices impact these services less than others? In a recent paper in Nature by David Crowder and colleagues, the question of how organic versus conventional farming affects predator and herbivore pathogen diversity and how this cascades to pest suppression. They show through a meta-analysis, that organic farms tend to support greater natural enemy evenness, and they hypothesize that greater evenness of enemies should better control pest populations, resulting in larger, more productive plants.

Picture from wikipedia

This result in itself is interesting, but they also carried out an elegant enclosure experiment where they manipulate the evenness of insect predators and pathogens and measure potato plant size. They found that even communities had the lowest herbivore densities and saw the greatest increases in plant biomass. Conversely, very uneven communities, typical of conventional farms, had the largest pest populations resulting in lower plant biomass accumulation.

While, multiple farming strategies are needed for adequate agricultural production, there are strong arguments for organic farms to be a important part of agricultural practice. These results show that organic farms have cascading effects on pest predators and pathogens and show that enemy evenness, as opposed to richness, has important ecosystem service consequences. To quote myself, evenness is a critical component of biodiversity, and much research has emphasized species richness, maybe at the detriment of studying evenness.

Crowder, D., Northfield, T., Strand, M., & Snyder, W. (2010). Organic agriculture promotes evenness and natural pest control Nature, 466 (7302), 109-112 DOI: 10.1038/nature09183

Teaching a quoll that cane toads are bad

Often, species become endangered because of multiple stressors, with habitat destruction taking the prize as the most egregious. However, often what pushes a species into extinction is not the main driver of endangerment. For example, passenger pigeon numbers were decimated by unabated hunting, but the proximate cause of extinction was likely an inability to thrive in low densities. Yet, seldom is the case where a known single species interaction is the primary cause of engangerment and maybe extinction. The northern quoll, Dasyurus hallucatus, is an endangered marsupial predator in Australia. The current major threat to the northern quoll is the invasion of toxic can toads. Quolls, being predators of small mammals, birds, reptiles and amphibians, readily attacks cane toads, which are toxic to quolls. Quoll populations have disappeared from areas invaded by cane toads, and extinction seems almost inevitable.

Given that the spread of cane toads into the remaining quoll habitats is inevitable, research, led by Stephanie O'donnell in Richard Shine's lab at the University of Sydney and published in the Journal of Applied Ecology, is underway to train quoll's to avoid cane toads. These researchers feed a subset of captive quolls dead toads laced with thiabendazole, a chemical that induces nausea. They then fitted individuals with radio collars and released these toad-smart quolls as well as toad naive ones. Some toad-naive quolls died quickly, after attacking cane toads. Only 58% of male naive quolls survived, while 88% of toad-smart males survived. While females seemed less likely to attack toads, 84% of naive females survived and 94% of toad-smart females survived!

See the video of a toad-smart quoll deciding not to eat a cane toad, its pretty cool.

O’Donnell, S., Webb, J., & Shine, R. (2010). Conditioned taste aversion enhances the survival of an endangered predator imperilled by a toxic invader Journal of Applied Ecology DOI: 10.1111/j.1365-2664.2010.01802.x

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.

Emergent linkages in seemingly unconnected food chains

Food webs are notoriously complex, and a difficult aspect of ecology is to offer a priori model-derived predictions of food web processes. There are some ecologists, such Neo Martinez and Jordi Bascompte, who have advanced our understanding of the general mechanisms of food web properties and dynamics through tools such as network theory. Such advanced approaches rely on direct interactions among species, or at least indirect interactions that are mediated through changes in abundance of different network players. However, what is missing from our general understanding of food web interactions is the role that behavioral responses can affect patterns of consumption and network connectivity.

Washington State University ecologists, Renée Prasad and William Snyder convincingly show how behavioral responses to predation can fundamentally alter food web interactions and link previously independent predator-prey interactions. They used two spatially independent insect predator-prey links in a novel, factorially-designed experiment. The two food chains consisted of a ground-based one, where ground beetles consume fly eggs and a plant-based one, where green peach aphids feed on the plants and are consumed by lady beetles. Under the ground-based chain only, the ground-based chain plus aphids, or ground-based chain plus lady beetles, the ground beetles consume a high proportion of the fly eggs. However, when both aphids and lady beetles are present, aphids respond to lady beetles by dropping off the plants and the ground beetles switch from consuming fly eggs to aphids. Under this last treatment, very few fly eggs are consumed, fundamentally altering the strength of the linkages in the two food chains and connecting them together.

This research highlights the inherent complexity in trying to understand multispecies systems, where the actors potentially have behavioral responses to other species, changing the nature of interactions. These types of responses may also generally increase the connectedness of such networks, which may result in more stable food webs, but this would need to be empirically tested. Regardless, this type of experiment offers food-for-thought to scientists trying to work general processes into a broad understanding of food web dynamics.

Prasad, R., & Snyder, W. (2009). A non-trophic interaction chain links predators in different spatial niches Oecologia DOI: 10.1007/s00442-009-1486-7