RESILIENCE AND STABILITY OF PERMO-TRIASSIC KAROO BASIN COMMUNITIES

Tags

biodiversity, extinction, food webs, modeling, paleo-food web, paleontology, simulations, trophic guild

Late Permian community dynamics

This is the second presentation that I made at the Annual Conference of the Geological Society of America in Vancouver last month. The presentation was part of a special session, “Extreme Environmental Conditions and Biotic Responses during the Permian-Triassic Boundary Crisis and Early Triassic Recovery”, co-organized my myself, Tom Algeo, Hugo Bucher and Arne Winguth. The session, spanning two days, was excellent, outstanding, and a lot of fun! I came away with the firm conviction that we are beginning to really understand the massive Permo-Triassic mass extinction, from its causes to consequences to recovery. It truly was a watershed “moment” in the history of the biosphere. The full program for both days can be found here and here. And, here is the abstract. An online copy of the presentation is available here.

RESILIENCE AND STABILITY OF PERMO-TRIASSIC KAROO BASIN COMMUNITIES: THE IMPORTANCE OF SPECIES RICHNESS AND FUNCTIONAL DIVERSITY TO ECOLOGICAL STABILITY AND ECOSYSTEM RECOVERY

ROOPNARINE, Peter, Invertebrate Zoology and Geology, California Academy of Sciences, 55 Music Concourse Dr, Golden Gate Park, San Francisco, CA 94118, proopnarine@calacademy.org and ANGIELCZYK, Kenneth D., Department of Geology, The Field Museum, 1400 South Lake Shore Drive, Chicago, IL 60605

A central question of the P/Tr extinction is the manner in which Permian ecological communities collapsed and E. Triassic ones were built. The end Permian Dicynodon Assemblage Zone (DAZ) has recently been resolved into 3 phases of the extinction spanning ~120ky, followed by the E. Triassic (Induan) Lystrosaurus Assemblage Zone (LAZ), offering an opportunity to examine the ecological dynamics of extinction and recovery in enhanced detail. We do this with 2 modelling approaches.

The first model assumes that populations exist in an energetic balance between consumption and predation. Communities are modelled as stochastic variants sampled from a space defined by species richness and functional diversity. Paleoenvironmental data from the DAZ indicate an increasingly seasonal, arid and drought-prone environment. The models were perturbed by simulated reductions of primary productivity. Results show that DAZ Phase 0 (Ph0) was a robust community resistant to low-moderate levels of perturbation with a well-defined collapse threshold. DAZ Ph1 and Ph2, however, exhibit highly variable responses and are significantly less resistant. LAZ similarly exhibits highly variable responses across minor variation of model configurations.

The second model assumes that communities are locally stable, i.e. minor perturbations are followed by asymptotic returns to equilibrium. During this return, however, communities can exhibit transient behavior during which perturbations can be greatly amplified. Amplification is likely to be important in unstable environments when the frequency of perturbations is shorter than the return time to equilibrium. Applying this model to DAZ and LAZ communities shows that the Karoo ecosystem became more limited in its responses to perturbation as the P/Tr boundary was approached, with Ph1 and Ph2 communities exhibiting very little transient behavior. LAZ in contrast exhibits increased transience.

The energetics and stability models are reconcilable in a history where the Karoo ecosystem became more ecologically stable as the extinction unfolded, yet more sensitive to cascading effects of species extinction and reductions of productivity. The Induan ecosystem was an unrecovered one, sensitive to both extinction and minor ecological disturbances.

Specimen collecting and cherry picking

Tags

biodiversity, california academy of sciences, endangered species, extinction, specimen collecting

In a recent opinion piece in Slate, Ben Minteer of Arizona State University continues to raise questions of the ethical legitimacy of collecting specimens for biological research. Minteer maintains that the risk to species, where population sizes might be small enough so that collecting represents a probabilistic extinction threat, outweighs the benefits to science and conservation. Unfortunately, Minteer is expressing an opinion, not the results of a carefully weighed and conducted analysis of data or facts. This is best highlighted by his example of the recent re-discovery of a species of New Guinea bat. Minteer states, “No scientist or conservationist today would deny the importance and value of describing a new species or confirming the return of one thought lost to extinction. But scientists also have a powerful ethical responsibility to minimize any and all adverse ecological impacts of their work.” Would that the world be so easily navigated. Today there are larger threats looming to biodiversity than at any time in the past 66 million years, and every one of those threats is the result of human actions. The threat of negative ecological impacts by scientists who are trying to document, explain and ultimately sustain what remains of the natural world pales hugely when compared to the threats of habitat destruction, the over-exploitation of species, and climate change. We will face very difficult decisions in the coming decades, and information is our friend, not our enemy.

Back to Minteer though. I think that his argument amounts to cherry picking and straw men. The reason for my position is best stated in a recent blog post by my colleague at the California Academy of Sciences, Dr. Jack Dumbacher. Jack explores the discovery of that very same bat picked by Minteer as an example, and he outlines very nicely the critical nature of the work. Please read his post. I’ll end here with an excerpt: “This study highlights the value of museum specimens in modern research, and the importance of taking specimens in modern field studies. Ironically, these studies were undertaken to assess the impacts of selective logging. The biggest threat to lowland forest in PNG is due to habitat loss from logging, mining, and oil palm conversion. One of the few things that might slow habitat loss is the fact that one little poorly known female bat was recently collected there.“

The legitimacy of collections for biological research

Tags

biodiversity, extinction

(copyright Python? via YouTube)

The April 18th issue of Science magazine included a piece by Ben Minteer of Arizona State University, and co-authors, “Avoiding (re)extinction“. The authors argued that the collection of specimens for biological research has, and may continue to place species at heightened risk of extinction, citing among other things stories such as the collection of the last remaining individuals of the Great Auk, as if those extinctions could be attributed to scientific collecting. The piece was very ill-conceived and poorly supported by evidence, basically constructing a straw man in the interest of argument (“But this is just contradiction.” “No it isn’t!”). Luiz Rocha, one of my colleagues here at the California Academy of Sciences, led a response which eventually involved 134 scientists hailing from 64 institutions around the world. Our response was published last week, also in Science. I cannot print any of the letters here, but I will include a link to our press summary.
SCIENTIFIC COLLECTIONS PLAY VITAL ROLE IN CONSERVATION BIOLOGY

Today is the Day of International Biodiversity

Tags

biodiversity, extinction, food webs

Today the UN and other organizations recognize the critical importance and threats to biodiversity around the world. The Species Alliance is recognizing the day by airing its documentary, Call of Life, on Free Speech TV (also streamed online). The documentary is followed by short interviews of myself (Peter Roopnarine), and Stuart Pimm. Please join if you can!

Science Today: Studying paleo-food webs

Tags

california academy of sciences, extinction, food webs, paleo-food web, paleontology

One of the most important ways species interact in an ecosystem? Food Webs. Learn how researchers study paleo and present-day food webs.
From the California Academy of Sciences.

Reining in the Red Queen

Tags

adaptation, evolution, extinction, paleontology, Red Queen

(The Victorian Web)

Geerat Vermeij and I just published a new paper in Paleobiology, entitled ” Reining in the Red Queen: The dynamics of adaptation and extinction re-examined.” The paper is partly a follow-up to my earlier paper on the Red Queens Hypothesis, reported previously in these posts (here and here), and partly the result of a discussion started between Vermeij and myself while attending the workshop that resulted in this paper on state transitions in the global biosphere. Here we argue that some of the fundamental assumptions of the hypothesis are flawed and that it therefore likely holds only under restricted circumstances. The full reference and abstract follow.

Vermeij, G. J. and P. D. Roopnarine. 2013. Reining in the Red Queen: The dynamics of adaptation and extinction re-examined. Paleobiology 39:560-575.

Abstract

One of the most enduring evolutionary metaphors is Van Valen’s (1973) Red Queen. According to this metaphor, as one species in a community adapts by becoming better able to acquire and defend resources, species with which it interacts are adversely affected. If those other species do not continuously adapt to compensate for this biotically caused deterioration, they will be driven to extinction. Continuous adaptation of all species in a community prevents any single species from gaining a long-term advantage; this amounts to the Red Queen running in place. We have critically examined the assumptions on which the Red Queen metaphor was founded. We argue that the Red Queen embodies three demonstrably false assumptions: (1) evolutionary adaptation is continuous; (2) organisms are important agents of extinction; and (3) evolution is a zero-sum process in which living things divide up an unchanging quantity of resources. Changes in the selective regime need not always elicit adaptation, because most organisms function adequately under many ‘‘suboptimal’’ conditions and often compensate by demonstrating adaptive flexibility. Likewise, ecosystems are organized in such a way that they tend to be robust and capable of absorbing invasions and extinctions, at least up to a point. With a simple evolutionary game involving three species, we show that Red Queen dynamics (continuous adaptation by all interacting species) apply in only a very small minority of possible outcomes. Importantly, cooperation and facilitation among species enable competitors to increase ecosystem productivity and therefore to enlarge the pool and turnover of resources. The Red Queen reigns only under a few unusual circumstances.

PNAS: Late Cretaceous restructuring of terrestrial communities facilitated the End-Cretaceous mass extinction in North America

Tags

Cretaceous, dinosaurs, extinction, food webs, mass extinction, modeling, networks, paleo-food web, paleontology, Robustness, Scientific models, simulations, Tipping point, trophic guild

That’s the title of our new paper, hot off the PNAS press. This study was a lot of fun, because it combines my food web work with one of the best known events in the fossil record. The lead author is Jonathan Mitchell, a graduate student at the University of Chicago. Jon became familiar with the food web work via Ken Angielczyk at the Field Museum, also in Chicago, a former post-doctoral researcher in my lab and close collaborator.  Jon wondered what Late Cretaceous, dinosaur-bearing communities would look like when subjected to CEG perturbations (just search this blog for info. on CEG!), and presented his results two years ago at the Annual Meeting of the Geological Society of America. The results were so intriguing that we decided then to explore the question in much greater detail, and ask what sorts of community and ecosystem changes unfolded in the years before the Chicxulub impact, and what role they might have played in the subsequent extinctions. And here are the results! I will list the full reference below, and you can obtain a complete copy of the paper from PNAS (sorry, not open access). Also, here are links to some news websites that have covered the paper, as well as the paper’s abstract. Enjoy!

EurekAlert, Science Daily, Science Codex

Jonathan S. Mitchell, Peter D. Roopnarine, and Kenneth D. Angielczyk. Late Cretaceous restructuring of terrestrial communities facilitated the End-Cretaceous mass extinction in North America. PNAS, October 29, 2012

ABSTRACT

The sudden environmental catastrophe in the wake of the end-
Cretaceous asteroid impact had drastic effects that rippled through
animal communities. To explore how these effects may have been
exacerbated by prior ecological changes, we used a food-web
model to simulate the effects of primary productivity disruptions,
such as those predicted to result from an asteroid impact, on ten
Campanian and seven Maastrichtian terrestrial localities in North
America. Our analysis documents that a shift in trophic structure
between Campanian and Maastrichtian communities in North
America led Maastrichtian communities to experience more second-
ary extinction at lower levels of primary production shutdown and
possess a lower collapse threshold than Campanian communities.
Of particular note is the fact that changes in dinosaur richness had
a negative impact on the robustness of Maastrichtian ecosystems
against environmental perturbations. Therefore, earlier ecological
restructuring may have exacerbated the impact and severity of the
end-Cretaceous extinction, at least in North America.

Extinction is not an Evolutionary imperative

Tags

extinction

Therefore, please stop using phrases such as, “Extinction is the fate of all species”. True, maybe 99.99% of all species that have ever existed are now extinct, but that is a frequency, not a law. Extinction results from a failure to acclimate on a short timescale, or to adapt on a longer, evolutionary timescales. The apparent extinction of species on the long run is the result of those failures, and is neither a requirement nor a certainty.

Number of predators per prey after extinction I: A start

Tags

extinction, food webs, link distribution, Network theory, networks

This series of posts are inspired by two questions that Jarrett Byrnes asked:

1. Given the extinction of E predators out of N, what is the probability that a prey species will still have at least one predator remaining?
2. Given E out of N, what then is the probability that all prey species will have at least one predator remaining?

As Jarrett and I have been discovering, these are actually quite difficult questions to answer in a general manner, i.e. for all topologies of a certain size!

Say we have a two trophic level food web with N predators, what is the probability that a prey species has at least one predator remaining after the extinction of E predators? The solution provided here depends on having the out-degree of prey species, and finding the probability that all predators of a prey species become extinct as a result of E. Say that the out-degree of the prey species is , then that probability is a hypergeometric solution

which reduces to

The probability then of a prey species having at least one prey is 1 minus the above

that is, the sum of the probabilities of having 1 predator, 2 predators, etc.

Example

Let the adjacency matrix of a food web be

where predators are rows and prey are columns. Our prey out-degree set is therefore {3, 2}. For E=1, both prey will have at least one predator since their out-degrees both exceed 1. For E=2, the possible resulting topologies are

For s=2

This is correct since our prey species of out-degree 2 (second column of A) has at least one predator in two of our three post-extinction topologies. The probability should be zero for s=3 (since E<s). If we add a third prey species, with s=1, making

then for E=1, the post-extinction topologies are

The probability that this third species has at least one prey is also 2/3.

A further example

So far so good, right? Well, Jarrett posed this example,

Notice that we now have two prey of out-degree 2. For E=1, the post-extinction topologies are

Applying the above formula yields

which is correct, since two of the three topologies maintain at least one predator for each prey. When E=2, the topologies become

Obviously, . But the formula gives

What went wrong?! The answer points to just how devilish the questions are, and how deceptive! There are two species of out-degree 2 (s=2) in the food web, hence the second term in the formula is squared (see above). BUT, the predator-prey topologies of the species are different, meaning that simple hypergeometric counting cannot work. We literally must list and examine all the post-extinction topologies, but this is prohibitively impractical for food webs and networks of even modest size (a dozen species). So there we stand. We currently have a partial solution, and I will explore the difficulty and the partial solution in the next post.

Instability in the Early Triassic!

Tags

competition, extinction, modeling, networks, paleo-food web, paleontology, Permian-Triassic, Scientific models

In a recent paper in the Royal Society Proceedings B, Randy Irmis and Jessica Whiteside verify a prediction of the CEG model regarding earliest Triassic terrestrial communities of the Karoo Basin in South Africa. Ken Angielczyk and I were interviewed by Wired Science for an article about the paper. Read it all here!

We predicted that communities in the Lystrosaurus Assemblage Zone would exhibit intrinsic instability in the face of even mild disruptions of primary productivity. More recently (and here), we explained that the intrinsic instability stemmed from the rapid diversification of small to medium-sized synapsid carnivores in the aftermath of the end-Permian mass extinction, coupled with very low species richness of herbivorous tetrapod prey, and the resulting intensity of competitive interactions among the carnivores. The recent Proceedings B paper seems to support our prediction on the basis of relative abundances of species of different trophic ecologies, characterizing those species as “boom and bust”. It’s always great to have model verification!

I think that there are some unresolved questions though:

1. We also suggested that one way out of the conundrum would be the increased specialization of the carnivores. Contrary to Irmis and Whiteside, I don’t agree that uneven relative abundances necessarily lead to demographic boom and bust cycles. Community dynamics are more nuanced and flexible than that.
2. The authors also point to probable environmental instability based on carbon cycles (measured as carbon stable isotope signatures). They valiantly overlap the short Karoo signature with the much longer and highly resolved marine signature. We simply have no good correlation of these signatures, and this is at least a nice attempt to highlight this ongoing issue.

Whether you can observe a thing or not depends on the theory which you use. (Einstein)