Monitoring northern biodiversity: picking the right trap for collecting beetles and spiders

Ecological monitoring is an important endeavour as we seek to understand the effects of environmental change on biodiversity. We need to benchmark the status of our fauna, and check-in on that fauna on a regular basis: in this way we can, for example, better understand how climate change might alter our earth systems. That’s kind of important.

A northern ground beetle, Elaphrus lapponicus. Photo by C. Ernst.

A northern ground beetle, Elaphrus lapponicus. Photo by C. Ernst.

With that backdrop, my lab was involved with a Northern Biodiversity Program a few years ago (a couple of related papers can be found here and here), with a goal of understanding the ecological structure of Arthropods of northern Canada. The project was meant to benchmark where we are now, and one outcome of the work is that we are able to think about a solid framework for ecological monitoring into the future.

A few weeks ago our group published a paper* on how to best monitor ground-dwelling beetles and spiders in northern Canada. The project resulted in over 30,000 beetles and spiders being collected, representing close to 800 species (that’s a LOT of diversity!). My former PhD student Crystal Ernst and MSc student Sarah Loboda looked at the relationship between the different traps we used for collecting these two taxa, to help provide guidelines for future ecological monitoring. For the project, we used both a traditional pitfall trap (essentially a white yogurt container stuck in the ground, with a roof/cover perched above it) and a yellow pan trap (a shallow yellow bowl, also sunk into the ground, but without a cover). Traps were placed in grids, in two different habitats (wet and “more wet”), across 12 sites spanning northern Canada, and in three major biomes (northern boreal, sub-Arctic, and Arctic).

Here’s a video showing pan traps being used in the tundra:

Both of the trap types we used are known to be great at collecting a range of taxa (including beetles and spiders), and since the project was meant to capture a wide array of critters, we used them both. Crystal, Sarah and I were curious whether, in retrospect, both traps were really necessary for beetles and spiders. Practically speaking, it was a lot of work to use multiple traps (and to process the samples afterwards), and we wanted to make recommendations for other researchers looking to monitor beetles and spiders in the north.

The story ends up being a bit complicated… In the high Arctic, if the goal is to best capture the diversity of beetles and spiders, sampling in multiple habitats is more important than using the two trap types. However, the results are different in the northern boreal sites: here, it’s important to have multiple trap types (i.e., the differences among traps were more noticeable) and the differences by habitat were less pronounced. Neither factor (trap type or habitat) was more important than the other when sampling in the subarctic. So, in hindsight, we can be very glad to have used both trap types! It was worth the effort, as characterizing the diversity of beetles and spiders depended on both sampling multiple habitats, and sampling with two trap types. There were enough differences to justify using two trap types, especially when sampling different habitats in different biomes. The interactions between trap types, habitats, and biomes was an unexpected yet important result.

Our results, however, are a little frustrating when thinking about recommendations for future monitoring. Using more than one trap type increases efforts, costs, and time, and these are always limited resources. We therefore recommend that future monitoring in the north, for beetles and spiders, could possibly be done with a trap that’s a mix between the two that we used: a yellow, roof-less pitfall trap. These traps would provide the best of both options: they are deeper than a pan trap (likely a good for collecting some Arthropods), but are yellow and without a cover (other features that are good for capturing many flying insects). These are actually very similar to a design that is already being used with a long-term ecological monitoring program in Greenland. We think they have it right**.

A yellow pitfall trap - the kind used in Greenland, and the one we recommend for future monitoring in Canada's Arctic.

A yellow pitfall trap – the kind used in Greenland, and the one we recommend for future monitoring in Canada’s Arctic.

In sum, this work is really a “methodological” study, which when viewed narrowly may not be that sexy. However, we are optimistic that this work will help guide future ecological monitoring programs in the north. We are faced with increased pressures on our environment, and a pressing need to effectively track these effects on our biodiversity. This requires sound methods that are feasible and provide us with a true picture of faunal diversity and community structure.

It looks to me like we can capture northern beetles and spiders quite efficiently with, um, yellow plastic beer cups. Cheers to that!


Ernst, C, S. Loboda and CM Buddle. 2015. Capturing Northern Biodiversity: diversity of arctic, subarctic and northern boreal beetles and spiders are affected by trap type and habitat. Insect Conservation and Diversity DOI: 10.1111/icad.12143


* The paper isn’t open access. One of the goals of this blog post is to share the results of this work even if everyone can’t access the paper directly. If you want a copy of the paper, please let me know and I’ll be happy to send it to you. I’m afraid I can’t publish all of our work in open access journals because I don’t have enough $ to afford high quality OA journals.

** The big caveat here is that a proper quantitative study that compares pan and and pitfall traps to the “yellow roof-less pitfall” traps is required. We believe it will be the best design, but belief does need to be backed up with data. Unfortunately these kind of trap-comparison papers aren’t usually high on the priority list.

Beetles from the North

I’m super-excited to announce new research from the lab, published yesterday with lead author Dr. Crystal Ernst.

Crystal’s paper focused on taxonomic and functional diversity of beetles across 12 sites in northern Canada, ranging from Labrador to the Yukon Territory, and from the bottom of James Bay all the way up to the tip of Ellesmere Island. This work is result of the Northern Biodiversity Program: a multi-institutional collaborative project about the ecological structure of northern Arthropods.

Crystal Ernst, on the tundra.

Crystal Ernst, on the tundra.

The paper was titled “Drivers and Patterns of Ground-Dwelling Beetle Biodiversity across Northern Canada” and in this research Crystal sorted and identified over 9,000 beetles from 464 species, and she classified the species by their functional ecology to assess how functional diversity may vary across the large spatial scale of this project. Instead of re-writing a summary here, I thought to use this blog post as an opportunity to reflect on what I see as the critical findings from this work, and why this is a paper that I’m incredible proud to be a part of.

  • To me, one of the more interesting findings of this work was that the functional diversity of beetles varied by latitude: although beetles do many things (e.g., herbivore, decomposers, carnivores), it doesn’t seem like all these functions happen at all latitudes. For example, although we document an impressive number of carnivores at all the sites, they are relatively more common in the more northern locations. This is a bit peculiar, and suggests that food-webs involving arthropods vary in some important ways depending on the biome. We also document that temperature is a major explanatory variable when considering functional diversity, which raises the important question about potential effects due to climate change. Indeed, should temperatures change in the north, this may affect the functional ecology of beetles, which in turn could affect other parts of the system.


Figure 1 from the paper: Fig 1. Map of the 12 study locations (North Pole Azimuthal projection), showing the spatial distribution of functional groups. These were pooled into trophic groups, and the pie charts show the proportion of the total site biomass represented by each trophic group

Figure 1 from the paper: Fig 1. Map of the 12 study locations, showing the spatial distribution of functional groups. These were pooled into trophic groups, and the pie charts show the proportion of the total site biomass represented by each trophic group

  • The research generally supported the well-known pattern in biogeography about how species richness decreases at more northern latitudes. When looking at which environmental variable may explain this pattern, temperature again came out on top. In other words, what beetles are found where is in part due to the temperatures in that region. Climate change scenarios therefore have significant potential effects on beetles in the north: beetles, like most other arthropods, are tightly linked to temperature. Even small changes in temperatures in the north may have big consequences for beetles.
  • One of the other big findings, to me, was the fundamental value of species-level data for an important taxa, across vast areas of Canada. Crystal recorded new Territorial and Provincial records for 15 beetle species, increasing knowledge about northern biodiversity. I’m also pleased that the data are fully available on-line, via Canadensys, so other researchers can access the information, re-analyze data, and benefit from and build upon this work.
  • The Arctic is special: it is a vast, cold, treeless landscape, with blankets of tundra, and permafrost underfoot. But it’s also special for beetles. After Crystal analyzed the community-level beetle data, using ordination methods, it became apparent that assemblages from the Arctic Islands of Canada were distinct from the sub-Arctic and north-Boreal sites. From a conservation perspective this is quite important. To some, the Arctic may come across as a big, ‘life-less’ region, with the odd polar bear roaming about, but in reality it hosts thousands of species, including hundreds of beetle species, and that beetle community is very different from what we find in other parts of North America. Special things deserve recognition and protection.
  • Every journalist I talked to has asked “Why beetles?” This is an easy one to answer: they fill virtually all roles in ecosystems, they are diverse, they are of interest to many people, and they are beautiful. The latter point is an important one, as it is important to capture curiosity and fascination about arthropods.


Carabus vietinghoffi. Photo by Henri Goulet.

A northern beetle: Carabus vietinghoffi. Photo by Henri Goulet.

In sum, this was a terrific project to be involved with, and our lab (and our collaborators) are thrilled that the efforts from the Northern Biodiversity program are showing up in the literature (for more examples, check out this, or this).

And rest assured, there’s more to come…

Meet the lab: Crystal Ernst

This is the first in a series of posts where each Arthropod Ecology lab member can introduce themselves. First up is PhD student Crystal Ernst:

I’m a Ph.D. candidate in the final stages of my program: these days I’m crunching out analyses and writing papers as I prepare to submit my thesis at the end of the term. As a community ecologist, I spend a lot of time thinking about how and why different species assemble together in space and time. These questions are foundational to the study of ecology and provide the overall framework for my research program, which uses beetles and other ground-dwelling arthropods to study the structure and determinants of terrestrial animal assemblages.

PhD student Crystal Ernst installing pan traps along the Dempster Highway (Yukon)

PhD student Crystal Ernst installing pan traps along the Dempster Highway (Yukon)

I have spent my summers conducting field research in gorgeous, remote regions of our northern territories, including Kugluktuk Nunavut and the Dempster Highway in the Yukon. My colleagues, members of the Northern Biodiversity Program, have contributed to the collection efforts as well, resulting in specimens being obtained from twelve different locations in the boreal forest, the subarctic and high arctic, spanning Canada coast to coast. I’m now neck-deep in the joy of interpreting the stories contained in my collection of specimens.

Specimens in pan trap (photo by C Ernst)

Specimens in pan trap (photo by C Ernst)


Sorting specimens back in the lab

Sorting specimens back in the lab

I’ve taken two approaches with this work. First, I’ve used a fairly traditional taxonomic approach to studying these animals: by identifying them morphologically (with a microscope and identification keys), I can associate each individual with a known insect species – although some new species have also been discovered! With this information I can describe the species richness (diversity) and distributions of different beetles in the north, and see which species are associated with each other at different northern locations. Secondly, I’ve looked at my arthropods from the perspective of their ecological functions – their roles in their environments. For example, some insects are responsible for pollenating plants, others are important decomposers, and others still are predators; arthropod assemblages can therefore be described in terms of the diversity and dominance of different functional groups. I am in the process of comparing taxonomic and functional assemblages found across northern Canada, and working to determine what aspects of their ecosystems (things like: temperature, wind, and sunlight; the diversity and structure of the plant community in which they live; soil characteristics) are associated with the way these assemblages are structured, and how they change over time and across space.

Three color morphs of Blethisa catenaria, a rare subarctic species (H. Goulet)

Three color morphs of Blethisa catenaria, a rare subarctic species (H. Goulet)

A fun complementary topic I’ve researched is the relationships between some high arctic ground beetles and a fascinating group of parasites called hairworms. I found a number of beetles from different locations to be infested with these worms; in one instance almost a quarter of the beetles were infected! The parasites are aquatic as adults and must first infect an aquatic insect (like a mosquito larva) before being transmitted to a terrestrial host (like a beetle) via the predation of the aquatic host by the terrestrial insect. To complete their life cycles, the worms somehow compel the beetles to enter the water, effectively forcing them to drown themselves so that the worms can emerge safely into their aquatic habitat. This discovery suggests an important link between the creatures living in terrestrial habitats and those in aquatic habitats and tells us about the arctic food chain: beetles must be eating mosquitos or other insects that have aquatic larval/immature stages. These prey items may, in fact, be a very important source of food. More work needs to be done to confirm this! In the meantime, I am excited to have found these associations – the fact that these particular species of beetles can be hosts for hairworm parasites is new information, and it appears that the parasite itself is a new species!

Pterostichus caribou with hairworms (C. Ernst)

Pterostichus caribou with hairworms (C. Ernst)

When I’m not writing my thesis or putting obscure little black beetles on pins, you can probably find me working at McGill’s Teaching and Learning Services, enjoying my time as a teaching assistant, networking on Twitter, mucking around in my vegetable garden (or putting said veggies in jars), walking my dogs, enjoying nature while canoe tripping with my partner, poking wildlife, or lifting heavy things at the gym. I’m on the hunt for a fantastic postdoctoral position that will allow me to continue studying different communities of living things in other ecosystems, and that factors that affect how they’re put together, and I’m excited about the many opportunities out there!

It’s a wrap! How about a thesis on Arctic spiders? How about two of them…?

This week I am thrilled to report that two of my MSc students have successfully completed their degrees! Both the projects are part of the collaborative Northern Biodiversity Program – a project aimed to quantify and understand ecological change with Arthropods from Canada’s north.

A BIG congratulations to Sarah Loboda and Katie Sim  – they are both tremendously talented students, excellent Arachnologists, and wonderful people to know.  Last night we had our annual Lab BBQ – and at that event, I was pleased to give Sarah and Katie a small token of appreciation.  Here’s a photo showing them both with their wolf spider photographs (photos by the incredible Thomas Shahan):

Katie Sim (l) and Sarah Loboda (r) - successful MSc students!

Katie Sim (left) and Sarah Loboda (right) – successful (& happy) MSc students!

Sarah Loboda’s thesis is titled Multi-scale patterns of ground-dwelling spider (Araneae) diversity in northern Canada. Her research focused on broad diversity patterns of ground-dwelling spiders collected from our 12 study sites, spread across Canada’s north. Our project spanned 30 degrees of latitude and 80 degrees of longitude –> yes that is a lot of land area! Sarah identified over 300 spider species from 14 families, and over 23,000 individuals.  Publications are forthcoming so I won’t give details here, except to say that we can learn a lot about diversity patterns over broad spatial scales using a study taxon such as spiders.

Here's where the Northern Biodiversity Program took our field teams!

Here’s where the Northern Biodiversity Program took our field teams.

Katie’s work (co-supervised by Prof. Terry Wheeler) had a different slant, but was still on Arctic spiders. Her thesis is titled:  Genetic analysis of Pardosa wolf spiders (Araneae: Lycosidae) across the northern Nearctic. The first part of Katie’s thesis was about understanding the phylogeographic history of the Arctic spider Pardosa glacialis, with particular attention to post-glacial dispersal patterns, as inferred by population genetics. The second part of her thesis was focused on whether or not there is enough evidence to suggest two northern Pardosa species should remain as separate species, or be merged into one – based on both molecular and morphological characters.  Let’s just say that Katie had to be a ‘field genius‘, ‘lab genius‘ and ‘spider genitalia genius‘.  Here’s an example of what she looked at, a lot:

The epigynum of a wolf spider species, (part of) the topic of Katie's research.

The epigynum of a wolf spider species, (part of) the topic of Katie’s research.

In sum, I am thrilled to see Sarah and Katie finish up their work, although their success also comes with a touch of sadness, as I will miss their daily presence in the laboratory.  Stay tuned… we shall soon report all the details from their research.

Seasonality of Arctic Beetles

I’m excited to report on paper written by Crystal Ernst, PhD student in my lab, and well known as the “Bug Geek“. This paper is a product of the Northern Biodiversity Program (yes, it sure is great that the papers from this project are starting to appear!), and will be one of Crystal’s PhD thesis chapters. The paper is titled Seasonal patterns in the structure of epigeic beetle (Coleoptera) assemblages in two subarctic habitats in Nunavut, Canada

A very nice Arctic beetle! (photo by C. Ernst, reproduced here with permission)

A very nice Arctic beetle! (photo by C. Ernst, reproduced here with permission)

Here’s a plain-language summary of the work:

Although we often think of Arctic systems as cold and lifeless, Canada’s tundra habitats are home to a high diversity of arthropods (insects, spiders and their relatives). Beetles are important insects on the tundra – filling ecological roles as predators (feeding on other insects), herbivores (feeding on plants), mycophages (feeding on fungi), and necrophages (feeding on dead or decaying animals). In this research, we wanted to find out what happens to ground-dwelling Arctic beetles as a function of seasonality. We were curious about whether different species occurred at different times during the short Arctic summer, and whether the functions of the beetles changes over the summer. This is an important area of study because beetles perform important ecological functions, and knowing how these functions change over time may have broader implications for northern ecosystems. This is especially relevant in the Arctic since these systems have a short ‘active season’, and climate change is disproportionally affecting northern latitudes. If climate change alters an already short summer, what might happen to the beetles?

This research was done as part of the Northern Biodiversity Program (NBP) – a broad, integrative project about the diversity of insects and spiders across northern Canada. The NBP involved collecting samples at 12 sites in the Arctic, but at one of these sites (Kugluktuk, in Nunavut) we had an opportunity to do a more detailed collection over the entire summer of 2010. This involved setting out traps for the entire active season, from June through to August. These traps were plastic containers sunk into the ground – beetles that wander along the tundra fall unawares into these traps, which contain preservatives, and are trapped until a researcher collects the samples. Traps were placed in wet and (relatively) dry habitats so that we could compare the two habitats. After the collections were returned to our laboratory, the beetles were identified to species, counted, and the biomass of the beetles was estimated – biomass lets us determine what happens to the ‘amount of beetles’ on the tundra in addition to figuring out ‘how many’ (abundance) and ‘what kind’ (species) were in the traps. The beetles were also classified into their key ecological roles. The data were then compared as a function of when traps were serviced to let us assess what happens to beetles as a function of seasonality.

We collected over 2500 beetles, representing 50 different species – remarkably, 17 of these species represented new Territorial records. This means that 17 of the species that were identified had never before been recorded in all of Nunavut! Although many ecological functions were represented by the beetles we collected, most were predators. We documented that wet habitats had different kinds of beetle species than the drier tundra habitats, even though the actual number of species between the habitats did not differ. We also uncovered a seasonal affect on the functions of beetles in the system – as the season progressed, the beetles tended to be represented more by predators compared to earlier in the season, which was dominated by beetles representing a diversity of functions. The mean daily temperature also related to the seasonal change that was observed in the beetles.

PhD student Crystal Ernst, happily working on the Arctic tundra.

PhD student Crystal Ernst, happily working on the Arctic tundra.

This work is one of the first to carefully quantify how beetles change during short Arctic summers. We found a diverse assemblage of beetles, filling a range of ecological roles. These ecological roles, however, do not stay the same all summer long, and the shifts in the beetles were related to mean daily temperature. Given that Arctic systems will be significantly affected by climate change, this is worrisome – if temperatures increase, or become more variable, this may affect ecosystem functions that are mediated by beetles. This is more evidence supporting the need to track climate change in the Arctic, and play close attention to the small animals of the tundra.


Ernst, C., & Buddle, C. (2013). Seasonal patterns in the structure of epigeic beetle (Coleoptera) assemblages in two subarctic habitats in Nunavut, Canada The Canadian Entomologist, 145 (02), 171-183 DOI: 10.4039/tce.2012.111

The greatness of pseudoscorpions

As you know, I’m quite passionate about Arachnology, from spiders, to harvestmen and Pseudoscorpions.  These are all some of the creatures that fall into the category of the ‘obscure and amazing‘.  On the topic of pseudoscorpions, a few very fun and interesting things have happened recently, and enough to warrant a short blog post.  I also promised that I would post a few more videos related to some research activities on the hunt for pseudoscorpions in the Yukon.

1. Just look at this SEM of a pseudoscorpion!

A little while ago, my Arachnid friends and colleagues from Alberta, Heather Proctor and Dave Walter, forwarded me a stunning image of a pseudoscorpion taken with a scanning electron microscope (SEM).  Dave was kind enough to give me permission to share it here:

SEM of a pseudoscorpion (Chernetidae) - copyright D. Walter (reproduced here with permission)

SEM of a pseudoscorpion (Chernetidae) – copyright D. Walter (reproduced here with permission)

There really is something lovely about getting up close and personal with these little Arachnids. I don’t know this species, but it’s definitely in the family Chernetidae – a relatively diverse family, quite common across Canada.  My favourite Yukon species, Wyochernes asiaticus, is also a Chernetid.   Dave Walter really does some magic with his SEM images, and you are encouraged to check out is macromite blog (his home bug garden blog is also worth a peek!).

2.  Just look at these videos about collecting pseudoscropions in the wild! 

Speaking of my favourite Yukon species, I took a lot of videos of field work in the Yukon last summer and I wanted to share a few with you, here.  Although our larger purpose for the trip was to complete some follow-up field work for the Northern Biodiversity Program, I also wanted to collect additional specimens of a wonderful pseudoscorpion species.  The first video provides some context to the work, and gives you a bit of a flavour of the landscape up near the Yukon – Northwest Territory border in Canada:

Typically, pseudoscorpions are not that commonly encountered.  In my experience, when they are encountered, you tend to see one or two.  What is truly amazing is the sheer abundance of this species found under rocks in creek/river beds in the Yukon.  Furthermore, you can see and collect multiple life stages, including females with eggs.  This short video gives a taste for this abundance.

The third and final video is a big goofy, and highlight the ‘collecting gear’ and appropriate field attire for becoming a “pseudoscorpion hunter“.  I am continually on a crusade to help generate enthusiasm for Arachnids, whether it is dispelling myths, or trying to inspire others to become Arachnologists (you know, we do need Arachnologists in Canada!).

One important caveat:  you may NOT simply run to the Yukon and flip rocks to collect pseudoscorpions – many parts of the world, including the Yukon, have strict guidelines about what you can collect.  Permits are required, and be sure to check into this before you plan on becoming an Arachnologist!

3.  Just look at this pseudoscorpion necklace!

To further illustrate my rather quirky obsession, I managed to find a wonderful person on Etsy who was able to make me a pendant with a pseudoscorpion design:

The pseudoscorpion necklace.  You want one.

The pseudoscorpion necklace. You want one.

Not only that, this design is actually from a photography I took a few years ago, and is an accurate depiction of the cosmopoliton species Chelifer cancroides.

Chelifer cancroides - my photo which was used to design the pendant

Chelifer cancroides – my photo which was used to design the pendant

I KNOW you want to get yourself one of these… start a conversation with Lynn.  Get yourself one of these necklaces and stand proud with other pseudoscorpionologists!

In sum, I do hope you find this post interesting, hopefully fun, and has whetted your appetite from more information about curious critters.

Stay tuned… I will continue to post more about Arachnids…

Assessing five decades of change in a high Arctic parasitoid community

As my colleague Terry Wheeler mentioned on his blog, our Northern Biodiversity Program team is thrilled to see post-doc Laura Timms‘s paper about Arctic parasitoid wasps published in Ecography!  Our team worked on Ellesmere Island, Nunavut, in 2010, and compared parasitoid wasps to historical collections from the same site that were made in 1961-65, 1980-82, and 1989-92. Parasitoid wasps are at the top of the insect food chain: they lay eggs inside or on top of other arthropods and the wasp larvae emerge after consuming their hosts – a gruesome but very common lifestyle for many types of wasps.  Species at higher trophic levels, such as these parasitoid wasps, are often the first to respond to new environmental pressures, including the climate change that is occurring rapidly in Arctic systems.

Laura identified a LOT of wasps, recorded the type of host attacked (e.g. plant-feeding hosts versus hosts that are predators), and the body size of two species of wasps that were commonly collected in all time periods.  We found no clear pattern of change in most aspects of the parasitoid wasp community on Ellesmere Island over past 50 years, even though temperature and precipitation have increased significantly during the same period. However, there were some signs that parasitoids of plant-feeding insects may be more affected more than other groups: one common parasitoid species that was abundant in 1960s hasn’t been collected since then, and the community in the 2010 study contained fewer parasitoids of plant-feeding insects than previous studies.

Screen shot 2013-01-28 at 10.52.40 AM

Some members of the Northern Biodiversity Program working in the Yukon in 2012. (l-r, Chris Buddle, Laura Timms, Crystal Ernst and Katie Sim)

Laura takes it as a good sign that no major changes in the ecology of the high Arctic parasitoid community have been observed, but isn’t taking it for granted that the community will remain unaffected for long.  At 82°N, Ellesmere Island is relatively isolated, but other research has found that parasitoid communities further south are changing dramatically (Fernandez-Triana et al 2011).

Laura has the following comment about our work: “We hope that our findings will be used as baseline data for ongoing monitoring on Ellesmere Island”, said Timms.  “We know so little about these high Arctic insect communities, we should learn as much as possible about them while they are still intact.


Timms, L., Bennett, A., Buddle, C., & Wheeler, T. (2013). Assessing five decades of change in a high Arctic parasitoid community Ecography DOI: 10.1111/j.1600-0587.2012.00278.x

Fernandez-Triana, J., Smith, M., Boudreault, C., Goulet, H., Hebert, P., Smith, A., & Roughley, R. (2011). A Poorly Known High-Latitude Parasitoid Wasp Community: Unexpected Diversity and Dramatic Changes through Time PLoS ONE, 6 (8) DOI: 10.1371/journal.pone.0023719