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Beneath Our Feet

Are there predators in Antarctic soils?

Map of Antarctica showing McMurdo Dry Valleys' location, created in Google Earth
with imagery from US Geological Survey and Landsat

By: Ashley Shaw, PhD student in the Graduate Degree Program in Ecology at Colorado State University. For her dissertation research, Ashley works with the NSF-funded McMurdo Dry Valleys Long Term Ecological Research program.


On first look, the McMurdo Dry Valleys of Antarctica are desolate. This cold desert is among the world’s most extreme environments and forms the largest ice-free area in Antarctica. It is frigid, dry, and windy. It’s dark there for half of the year. During the austral summer, the one long day when the sun shines, the soils thaw and refreeze as the sun warms dark soils despite temperatures remaining below freezing. The UV levels are high (thanks, ozone hole!) and a constant threat due to 24 hours of daylight. Despite these challenges, there is life in the dry valleys.

In the dry valleys, the largest terrestrial animals are soil invertebrates: tardigrades, nematodes, and rotifers, along with a few springtails and mites. The soils they inhabit have their own suite of harsh factors and often have high pH, salt, and nitrogen due to deposition and build up that is not flushed out by precipitation. Not only is there infrequent snow, there is little liquid water available at all. No vascular plants survive here, and there are only trace amounts of organic carbon in the soils. Until now, environmental factors such as temperature, pH, salinity, and water availability were considered to be the only drivers of soil invertebrate communities.

Soil sample collected in Taylor Valley, Antarctica.
Photo by A. Shaw

While one tough nematode, Scottnema lindsayae, dominates and thrives in the dry soil that makes up >95% of the landscape, most of the soil invertebrate biodiversity prefers the scarce wetted margins of streams and lakes. These areas often have cyanobacterial mats – orange, black, and green - that sustain the soil food webs. In and around these mats, nematodes, rotifers, and tardigrades prosper. These animals share a habitat and are regularly extracted from the same soils. Could their interactions – such as predator-prey or competition – shape the communities in these soils?

Eudorylaimus antarcticus extracted from Taylor Valley soil
as seen under the microscope.
Photo by A. Shaw

The nematode Eudorylaimus antarcticus prefers wet soils, but is sometimes also found in dry soils with S. lindsayae. E. antarcticus is member of the order Dorylaimida, which contains many omnivore-predator nematode species. Given its close relationship to other predators, E. antarcticus could be a potential predator in the dry valleys. However, previous studies have only found evidence that it eats algae. We set out to test whether E. antarcticus could be a predator in the dry valley soils.

Stable isotopes are a great approach for understanding food webs. They are similar to radioisotopes (but have the benefit of not being radioactive!) and are also traceable. We can understand what an organism’s diet is like, because it picks up a detectable signature from its food sources. We used stable isotopes to test each invertebrate group’s position in the soil food web. To do this, we collected soil from both wet and dry habitats in the dry valleys. We extracted invertebrates from the soil, identified them to species under a microscope, and then collected (by hand, using an eyelash tool) hundreds of individuals by taxa. These invertebrates were analyzed for their 13C and 15N isotopes.

The isotope results showed that rotifers, tardigrades, and several nematode species were grazers – eating bacteria and algae. But E. antarcticus was different: its isotopes put it the top of the food web. This new evidence, combined with previous evidence that it eats algae, showed that E. antarcticus is an omnivore-predator.

This is big news for cold desert ecology. Not only is this the first solid evidence of a nematode predator present, it also opens the door to testing how biotic interactions such as predator-prey relationships might shape communities in these soils. This is especially important because the landscape is predicted to become wetter and more connected in the future. Thus, species ranges may shift, altering interactions or creating new contacts between species. Understanding species’ relationships to one another is the first step in predicting how their interactions might shape soil communities.

Antarctic soils are far from desolate. Life abounds in these soils, where there are still many questions to be explored!

View from helicopter en route to our field site.
Photo by A. Shaw

Lake Bonney, Taylor Valley, Photo by A. Shaw


This work was published recently in the journal, Polar Biology, and can be found here:   Tags: 

Lifestyles of arbuscular mycorrhizal fungi

Examples of AM fungi hyphae (a, c) and
vesicles (b, d) in corn (a, b) and prairie roots (c, d).
Image from Bach et al. 2018, Ecology

By: Elizabeth Bach, Executive Director, Global Soil Biodiversity Initiative, School of Global Environmental Sustainability, Colorado State University


One of my first experiences with soil ecology was a summer Research Experience for Undergraduates, during which I got to spend a lot of time looking at arbuscular mycorrhizal fungi within roots. I had never really observed these microscopic relationships before, and seeing the ways the fungal bodies lived within the roots amazed me.  Hyphae extended out like fingers from the root, searching the soil for nutrients and water.  Vesicles appeared as small spheres, full of lipids that the fungus could consume for energy when the host plant quit delivering carbohydrates.

Fast-forward six years. I still loved working with soil microorganism and was a few years into a PhD program.  I had the opportunity to introduce this exciting microscopic world of mycorrhizal fungi to a couple of undergraduate assistants.  Over the course of several weeks of counting arbuscular mycorrhizal fungi on roots from corn (maize) and tallgrass prairie plants, these students, Giselle and Kira, noticed that the corn roots had more hyphae than the prairie roots and the prairie roots had more vesicles.  They asked why, and I didn’t know the answer, so we went looking in the scientific literature. Interestingly, we didn’t find many previous studies that could explain the pattern.

Giselle and Kira went back and specifically counted the rate of hyphae and vesicle presence on the roots, and it turned out corn roots had 6 times greater hyphal presence rate and prairie roots had 3 times greater rate of vesicle presence!  It also turned out that the rate of presence of these structures varied across the growing season, with hyphae colonization peaking in August for all roots, and vesicle presence peaking in July, for the prairie roots, and September, for corn roots.  Giselle and Kira shared their findings with poster presentations at our university (Iowa State University) and at national scientific meetings (American Geophysical Union and Ecological Society of America).

This data story remained interesting, but didn’t seem like enough to pursue a classic scientific publication. In 2017, the journal Ecology rolled out a new article type: The Scientific Naturalist.  The section is dedicated to short essays, sharing an observation of the natural world that captured the wonder of ecological discovery and could lead to new hypotheses and deeper research.  Immediately, Giselle and Kira’s work came to mind and I reached out to them to ask if they might be interested in working with me to write up their observations.  Over the next few months we, along with our lab group’s PI Kirsten Hofmockel and fellow AM fungi researcher Jonathan Bauer, worked to write-up our story to share.  The dynamic life of arbuscular mycorrhizal fungal symbionts was published January 24, 2018. It has been a delight to work with and learn from scientists from across the spectrum of experience on this project, and we are excited to see what future work may grow from this work!


Elizabeth Bach is Executive Director of the Global Soil Biodiversity Initiative housed in the School of Global Environmental Sustainability at Colorado State University.

Giselle Narvaze-Rivera is a Master’s of Science student in Physical and Biological Anthropology at Iowa State University.

Kira Murray is a consulting geologist at Freestone Environmental, Richland, WA.

Jonathan Bauer is a postdoctoral researcher at Michigan State University

Kirsten Hofmockel is Lead Scientist for Integrative Research at the Environmental Molecular Science Laboratory at Pacific Northwest National Laboratory.


Original Article:

Bach, E.M.; Narvaez-Rivera, G.; Murray, K.; Bauer, J.T.; Hofmockel, K.S. 2018. The dynamic life of arbuscular mycorrhizal fungal symbionts. Ecology doi:10.1002/ecy.2096


Bridging the gap in freshwater global warming research

An example of sediment sampling from streams
used to characterise and quantify microbial
taxonomic and functional diversity
Image: H. Prentice

Dr Kate Randall, Research Associate and Molecular Microbial Ecologist at the University of Essex, UK is part of a multidisciplinary research team investigating the impacts of temperature on freshwater ecosystems as part of a ‘Ring of Fire’ NERC-funded research project led by Professor Guy Woodward from Imperial College London, UK.

Like soil, marine and freshwater sediment are places where organic and inorganic material is deposited, stored, or used as nutrients and energy to support the main hub of biodiversity within these environments. Due to the isolated locality, influx of water and contents, freshwater ecosystems are particularly vulnerable to changes in land use, biotic and abiotic conditions. The effects of which are manifesting as faster rates of biodiversity loss compared to terrestrial ecosystems (Dudgeon et al. 2006).

The majority of us are aware of the unprecedented rates of warming our planet is experiencing, but the effects on freshwater systems in particular sediment biodiversity and functioning are not well understood. Most research to date focuses on the responses of a small range of larger-bodied organisms to warming. In reality an understanding of foodwebs, combined with measurements of multiple process rates (i.e. denitrification, methanogenesis) is required to create ecological networks that refine understanding of freshwater responses to warming. In addition, the microbiota are largely underrepresented in the multitrophic climate change studies that do exist (Sarmento et al. 2010), especially within freshwater environments. Given that the ubiquity and diversity of microorganisms is considered to be the greatest on the planet, with key roles in biogeochemical nutrient cycles (Krumins et al. 2013), this needed to be addressed.

Through the collection of field samples, mesocosm and laboratory microcosm experiments, the ‘Ring of Fire’ project has embarked upon bridging the gaps in freshwater responses to global warming by conducting multidisciplinary research that spans multiple spatial scales and adopts a unique genes to ecosystems approach detailed below. Collectively, these components of the ‘Ring of Fire’ project will offer enough data to allow computational approaches such as machine-learning and ensemble species-distribution models to characterise and refine predictions of species and community distributions, food web structure and dynamics responses to warming across spatial and temporal scales, at a level previously unexplored within these vulnerable and highly valuable ecosystems.

Large spatial scale field based sampling of freshwater streams along natural geothermal gradients

Two Icelandic streams – one is 5 ºC and the other
is 20 °C and are only meters apart
Image: M. Jackson

Geothermal activity provides an opportunity to study long-term warming effects on natural freshwater streams as the activity generates a temperature gradient along which streams are located. Our large scale field research targets geothermal activity in multiple high latitude regions, within which, temperature does not correlate with other physicochemical stream variables. This component of the project will provide knowledge of how freshwater streams in an area experiencing some of the fastest rates of warming are responding, and importantly, act as indicators for freshwaters elsewhere in the future (O’Gorman et al. 2014). A core team of us were tasked with venturing to 5 sites during summer 2016 and 2017 (Iceland, Alaska, Greenland, Svalbard and Russia), where stream temperatures ranged from 2-35 oC. The multidisciplinary team is divided into molecular microbial ecologists, freshwater ecologists and biogeochemists who have successfully collected samples from stream sediment, the water column and biofilm. Careful planning developed a sampling protocol that was consistent across all streams, protected the integrity of samples for each research group, and allows complementary downstream data collation.

Global mesocosm warming experiments (Mediterranean to the Arctic)

A number of large mesocosm ponds have/are being established across the globe such as the UK (, Iberia ( and Denmark to investigate warming effects within different climatic zones. Here ponds can be heated at different temperatures and combined with additional stressors associated with climate change, such as spiked warming events, drought regimes and variations in nutrient availabilities. This will allow regulation of treatments and detection of causal relationships between these variables that would not be possible in the field.

Initial 96 mesocosm pond set-up at
Imperial College London – Silwood campus

Hamilton StarLet - Automated robotic liquid handling
to prepare mock microbial communities and to assist
sample preparation for molecular microbial work

Lab based microbial microcosm experiments – Ecological and evolutionary responses to warming

Environmental samples collected from the geothermal streams and global mesocosm sampling will be used to isolate and seed into various combinations. These mock communities will then be subjected to changes in temperature and nutrient availability to disentangle the effects on complex networks of interacting microbial groups for which mechanistic drivers could not be determined within the natural environment.

Collectively, these components of the ‘Ring of Fire’ project will generate enough data to allow computational approaches such as machine-learning and species-distribution models to characterise and refine predictions of species and community distributions and high resolution ecological networks across spatial and temporal scales at a level previously unexplored within these vulnerable and highly valuable ecosystems.


Further Reading

Dudgeon, D., Arthington, A.H., Gessber, M.O., Kawabata, Z., Knowler, D.J., Leveque, C., Naiman, R.J., Prieur-Richard, A., Soto, D., Stiassny, M.L.J., Sullivan, C.A., 2006. Freshwater biodiversity: importance, threats, status and conservation challenges. Biol. Rev. 81, 163-182. doi:10.1017/S1464793105006950

Krumins, J.A., van Oevelen, D., Bezemer, T.M., De Deyn, G.B., Gera Hol, W.H., van Donk, E., De Boer, W., De Ruiter, P.C., Middelburg, J.J., Monroy, F., Soetaert, K., Thebault, E., van de Koppel, J.A, Viketoft, M., van der Putten, W.H., 2013. Soil and Freshwater and Marine Sediment Food Webs: Their Structure and Function. 63, 35-42. doi:10.1525/bio.2013.63.1.8

O’Gorman, E.J., Benstead, J., Cross, W.F., Friberg, N., Hood, J.M., Johnson, P.W., Sigurdsson, B., Woodward, G., Climate change and geothermal ecosystems: natural laboratories, sentinel systems, and future refugia. Glob Change Biol. 20, 3291-3299. doi:10.1111/gcb.12602


Major threats to soil ecosystems from a combination of invasive species and climate change

An Isotomurus springtail species. Alien species
like these have, on average, greater resistance
to high temperatures than their indigenous counterparts.
Image: ChownLab, Monash University.

By Charlene Janion-Scheepers, Monash University, Australia


A study examining heat tolerance in alien and indigenous springtails, key soil arthropods that effect many aspects of ecosystem functioning, finds that tolerance of warming, such as that associated with climate change, is on average much more pronounced in the alien species than their indigenous counterparts, with little scope for adjustment by evolutionary change or phenotypic plasticity, suggesting that the impacts of biological invasions on soil systems will be exacerbated by climate change.


Soil ecosystems are critical for agriculture, biodiversity and human well-being. Poor soil health means a poor planetary outlook. A study published in Proceedings of the National Academy of Sciences of the USA by a Monash University team shows that a new threat faces soil sustainability everywhere. The team found that, from the polar regions to the tropics, invasive soil-dwelling species are typically better able to cope with warming than their indigenous counterparts. Climate change will benefit invasive species, suggesting major changes to the functioning of ecosystems, and potentially bad news for the United Nation’s Sustainable Development Goals of conserving and restoring terrestrial ecosystems (SDG 15: Life on Land), and ending hunger and malnutrition (SDG 2: Zero Hunger).

The work was conducted on springtails – small soil invertebrates that are globally ubiquitous, and influence both soils and the aboveground ecosystems which rely on them. Together with other invertebrates, such as earthworms, these animals make soil ecosystems work. As with earthworms, humans have moved springtail species unintentionally around the world from their natural homes to new environments. What the study shows is that these alien species are much more tolerant of high temperatures than their indigenous relatives.

Springtails, such as this neanurid species,
play a key role in terrestrial ecosystems.
Image: ChownLab, Monash University.

"The impacts of soil invasives are becoming increasingly well known” said Dr Charlene Janion-Scheepers, lead author, “What we have found is that alien invasive species will thrive under climate change, much more so on average than local species, irrespective of where one looks”.

Co-author, Associate Professor Carla Sgrò, further added “We also tested to see if indigenous local species might evolve greater tolerance. They simply cannot. So, for the foreseeable future, we will be in a world where on average, invasive species are going to be the winners.”

Mounting concerns about the impacts of biological invasions for agriculture and terrestrial ecosystems have recently made world headlines. What this study does is demonstrate that impacts will be exacerbated by changing climates.

“The Paris Agreement on reducing emissions to combat climate change becomes all the more important when we see results like this” emphasized team leader Professor Steven Chown. “What’s at stake is nothing less than the health of our ecosystems.”


Primary publication: Janion-Scheepers et al. 2018. Basal resistance enhances warming tolerance of alien over indigenous species across latitude. Proceedings of the National Academy of Sciences of the USA 115, 145-150,



How much soil is lost every year?

By Panos Panagos and Cristiano Ballabio European Commission’s Joint Research Centre, Directorate D – Sustainable Resources, Land Resources Unit, Ispra, Italy

Amount of soil lost by water erosion in 2012 at 250 × 250 m resolution for 202 countries (approx. 125 million km2).
Image from Borrelli et al. 2017

Soil is home of thousands of organisms, targeted by hundreds of soil biology and ecology studies. Soil is not an infinite resource, therefore, as soil biologists/ecologists, the raw material behind all of your findings is eroding away. The link between soil erosion and biology/ecology is stronger than what you may expect. As soil erosion modelers, our work improves estimates of soil loss in order to clearly assess its effects on soil life and, eventually, to develop actions to control it.

The question is simple: how much soil is lost every year on Earth due to erosion? Human activity and changes in land use lead to increased soil loss, which in turn degrades nature's nutrient cycling system, diminishes land productivity and affects the capability of soil to act as a habitat. Furthermore, erosion is becoming more and more an issue due to intensive meteorological events associated to climate change. According to the Status of the World's Soil Resources, recently published by the Food and Agriculture Organization, soil erosion is the main threat to soil (and soil dwelling organisms) worldwide. Still, the precise amount of soil lost annually remains unclear.

So far, a reliable assessment of soil erosion at global scale was missing. In December 2017, this gap has been partially filled by our new study. The analysis, produced by a group of researchers led by the European Commission's Joint Research Centre, the University of Basel and the Centre for Ecology & Hydrology, was published in Nature Communications: An assessment of the global impact of 21st century land use change on soil erosion.

We have developed an unprecedentedly high resolution (250 × 250 m) global potential soil erosion model, using a combination of remote sensing, GIS modelling and census data. The model result, an estimated 36 billion tons of soil eroded per year, is at least two times lower than previous annual soil erosion reference values.

As for biodiversity, we identified hotspots where erosion is more intense. The greatest amount of soil loss is estimated in Sub-Saharan Africa, South America and Southeast Asia. This means that some of the countries with less developed economies were estimated to experience the highest soil erosion rates. In particular, some of the largest and most intensively eroded regions are in African equatorial area (0.26 million km2, 3.2% of the region). Moreover, we calculated the spatial and temporal effects of land use change between 2001 and 2012. Our findings indicate a potential overall increase in global soil erosion mainly driven by cropland expansion (+0.22 million km2) and forest decline (−1.65 million km2).

Finally, we also evaluated the potential offset of the global application of conservation practices in agriculture. Our study estimated that, if applied correctly, conservation practices could save over a billion tons of soil per year. The highest reductions in soil loss due to conservation agriculture were estimated to occur in South America (16%), Oceania (15.4%) and North America (12.5%). The message is clear: soil erosion can be reduced if soil conservation practices are adopted in agriculture.

Overall, our research shows how soil erosion is a hot issue that requires actions to curb it, especially as it greatly affects developing countries. We hope that our work will promote the inclusion of soil erosion as one of the priorities in the environmental political agenda. This work will also help soil biologist and ecologists more aware of soil erosion and the impact it can have on their subject of study. Thus, we look forward to future studies integrating soil erosion modelling and soil biology and ecology. Indeed, another question arises: how do soil organisms affect the amount of soil eroded annually? It is up to soil biodiversity scientists to give an answer to that.



Digging for data gold in soil

By Kathe Todd-Brown, USA @KatheMathBio


I’ll frequently hear my field and lab colleagues disparaging their data sets: “Oh this core is constantly giving high values for no reason!” or “We just couldn’t get any methane out of this core, and then we found a rock.” As a researcher, you know all the warts of your own data set and it’s sometimes hard to see the gold underneath.

And your data set is gold. The time, money, and general effort invested in both designing the experiment/survey and then collecting the data to capture a single point in time that will never be exactly replicated again, all makes data incredibly valuable. In a very real sense, scientific data are the Truth that we are trying to understand as scientists. Especially when viewed in aggregation with other data to exact general scientific insight.

Data deserves respect, both yours and our colleagues. This is already common practice with field and lab notebooks. Who hasn’t opened a senior mentors old field notebook with a certain amount of awe and reverence to look at sketches of soil profiles or coffee stains over detailed protocol notes. But all too often researchers see publication as the end of the road for their data. While many data don’t even make it this far (publication bias is another post), this shortchanges the enormous potential for re-use and additional insights that could be gained from most data sets.

It goes without saying that soils are incredibly heterogeneous. Move over 10 centimeters, resample, and you get huge variations in your soil measurements. This makes large data sets critical for extrapolating generalizable insights. But most soil measurements are laborious, limiting most sample sizes to such small numbers that statisticians typically throw up their hands in despair. Individual studies will try to get around this by restricting the scope of their conclusions, homogenizing soil samples, or other methods. But another way around this is to pool data sets after the original study is concluded.

The more data you have, the more valuable it is. But there is a lot of work to get to a multi-data set harmonized data base. There are several hurdles to data re-use but they loosely fall into availability, discoverability and harmonization.

Is the data available? Data locked in basement filing cabinets is literally inaccessible unless you happen to have that key. Increasingly ‘contact the PI’ is an inadequate data policy for many funding agencies and academic journals. And for good reason. Individual researchers move around, leave the field entirely, or simply lose data. Making your data available through a university library, society archive, or some other long term institute is critical for preservation. But it is only the first step.

Once that data set is archived it needs to be discoverable. In the age of Google it still surprises many people how hard it is to find data once it is archived. Since data sets typically come in many different formats, data sets are not indexed directly on the data itself but on meta-data provided with the archived submission. This meta-data description is thus critical and frequently impossible to standardize. If you are investigating a new phenomenon there may be no standard way to describe this in the meta-data at the time of submission. In my opinion, the associated manuscripts makes the best advertisement for an archived data set. So it is important to link that manuscript to the archived data at publication. But there is active research in semantics and around dynamically developing control vocabularies to try to solve this problem.

Finally, if you manage to get the data out of basement storage and adequately described, there remains the third hurdle: harmonization. Is there enough information about the measurements and methods to be able to intelligently compare the data to other data sets? As mentioned previously, data sets are frequently in unique formats and may have new or otherwise distinct measurement protocols that makes automatic data ingestion intractable. Harmonizing the data set to make it comparable to a broader data collection requires expert understanding of the context of the data.

This final harmonization effort can be particularly tricky because you need to maintain a direct link to the original data set. Much like a field notebook or set of lab protocols, a script or computer program robustly preserves data provenance. Manual entry, transcribing data from one templet to another, is generally error prone and can be difficult to reproduce when reviewing the resulting meta-analysis. Sometimes manual transcription can’t be avoided but hand crafting unique scripts to process individual datasets provides the best way to harmonize data sets. Scripting data translation instead of manual entry is both explicit and reproducible, providing a clear line of data provenance from the original data file to the final data product.

All three stages -archiving, discovery, and harmonization- are laborious. Finding a repository to take the data, adequately describing what is there in the meta-data, and writing harmonization scripts is not glamourous work. No one will win a Nobel for their beautifully complete meta-data, but it is necessary.

Global soil maps, for example, can only be constructed by combining the results from soil surveys of different nations and regions. These maps are critical to benchmarking the land-carbon cycle of Earth system models used to inform anthropogenic emissions targets. The land-carbon models themselves rely on other data collections for parameterization and validation during model development. If you want your data to be of broader use to the community and live on beyond an individual project, it has to be preserved intelligently.

Data is scientific gold whose value increases as more data is collected. This is particularly true for a highly heterogeneous system like soils. Contributing data to the broader scientific collection requires that data be available, discoverable, and harmonize-able. While meeting these requirements can be laborious, the long-term rewards to the broader community are significant.


Kathe Todd-Brown is a computational biogeochemist at the Pacific Northwest National Laboratory in Richland, Washington. She initially trained as a mathematician but it was a bit too clean. She transitioned to soil carbon cycling and couldn’t be happier to work in one of the most interesting systems on the planet.


Soil biodiversity at continental scale, a call for support and collaboration

By Alberto Orgiazzi, European Commission’s Joint Research Centre, Directorate D – Sustainable Resources, Land Resources Unit, Ispra, Italy

Over the past 10 years, the European Commission’s Joint Research Centre (JRC), base in Ispra, Italy (nearby Lake Maggiore… a great place to work at) has been responsible for the organisation of the largest soil survey ever to be carried out across Europe. Named the LUCAS (Land Use/Land Cover Area Frame Survey) Soil, this campaign kicked off in 2009, with 19,000 samples being collected from across Europe. A second round of sampling was carried out in 2015 from about 23,000 points all over the continent, the soil samples of which are now being physically and chemically analysed. The results of these initial surveys are summarized in a new review paper: LUCAS Soil, the largest expandable soil dataset for Europe.

1,000 locations (and their land cover type) across Europe where soil samples will be collected,
as part of LUCAS Soil survey, for soil biodiversity assessment in 2018. Image from Orgiazzi et al. 2017

So far, the focus of LUCAS was exclusively on measuring soil properties such as texture, cation exchange capacity, pH, organic carbon, nitrogen and many others. One element was missing from this picture: the ‘alive’ component of soil, the soil biodiversity. The good news is that this element will be included in the third round of LUCAS, making it the first ever pan-European assessment of soil biodiversity.

Thanks partly to the success of the first ever Global Soil Biodiversity Atlas, which is demonstrating the increasing interest in soil life not only among scientists but also among policy makers, the JRC plans to analyse the biodiversity of soil organisms about 1000 of the total 26,000 points by means of DNA-fingerprinting techniques. The second piece of good news, at least to me, is that I am the postdoctoral researcher responsible for all that. I am very excited at this opportunity, and when they offered it to me I immediately said: I’m in. However, I admit that I am also pretty scared. Of course, I am not alone in this effort, and my colleagues are already helping me, but still I have been the one who drafted the protocols for sampling, storing and shipping the soil samples, and selected the organisms to be considered. This is likely one of the most challenging and, at the same time, fascinating tasks of my life, at least from a professional point of view.

Would you like to put yourself in my shoes? I have had a year and a half to organise everything, from the selection of sampling points to the choice of protocols for DNA extraction and amplification. So, I spent the last few months thinking about possible ways to plan the future work. Then I got a brainwave. Why not turn this responsibility into an opportunity? Now I have in my hands the possibility to contribute the European part of this global map/assessment of soil biodiversity. And I do not want to do this alone.

I am already in touch with people in Australia and Africa that are studying the distribution of soil life through DNA tools (Bissett et al., 2016; African Soil Microbiology Project). However, I am sure there are several other research groups worldwide that are planning to do something similar in the near future, for example in the frame of the recently proposed National Microbiome Project in the US and the China Soil Microbiome Initiative in China. And, perhaps, you are also planning some sampling of soil biodiversity. Those are the people I am addressing; let’s bring our ideas and efforts together in order to develop a common strategy. That is the purpose of the call that we have recently launched in our review (Orgiazzi et al., 2017). I am issuing this public call in order to reach the broadest possible audience.

You can contribute in many ways. You can propose possible strategies for large-scale DNA fingerprinting of soil organisms (see our review for details). And you can also actively contribute to both LUCAS Soil and global soil biodiversity assessment. In 2018, you may consider the possibility to collect some soil samples (not just across Europe) and analyse them through our protocols (by the way, we decided to use those proposed by the Earth Microbiome Project – get in touch for details). In this way, you will have your own LUCAS Soil points. That will make you able to compare your data to ours (1,000 samples… not bad) and address ecological/biological questions of your interest. At the same time, you will help out with the development of the first global soil biodiversity maps. Last but not least, you may also consider the possibility to visit one or more LUCAS points, collect your own samples and make whatever additional analysis (e.g. mesofauna and earthworm sampling). Then, we can combine our data. What more could anyone wish for?

I really look forward to receiving your inputs. My email is

Some of the material (1,000 polystyrene boxes and 4,000 ice blocks) that will be used to collect
fresh soil samples for soil biodiversity analysis. Image by A. Orgiazzi


Bissett A, et al. (2016) Introducing BASE — the biomes of Australian soil environments soil microbial diversity database. Giga Sci, 5: 21.

Orgiazzi A, et al. (2017) LUCAS Soil, the largest expandable soil dataset for Europe: a review. Eur J Soil Sci, DOI: 10.1111/ejss.12499


Soil Carbon Modelling with Soil Fauna and Humus Forms

An example of a mull humus form.
Image by D.Hoffman

By Cindy H. Shaw, Oleg Chertov, and Darrell Hoffman*

Soil fauna are key agents in different types of organic debris processing in the forest floor and mineral topsoil, resulting in humus forms. The humus form that develops at any given site is the result of the amounts and types of litter inputs (such as wood, leaves, roots) from local vegetation, temperature and moisture conditions, and the dynamics of soil biota including fauna, fungi, and bacteria. Humus forms are most broadly divided into three orders: mor, moder, and mull. Each order is associated with a set of physical properties that are easily observed and are related to processes occurring in the forest floor and the underlying mineral soil.

An example of a moder humus form.
Image by C.McNalty

Mors are characterized by a distinct boundary between organic and mineral soil horizons, often with plentiful plant roots and fungal hyphae throughout the organic layers. Mulls develop when there is sufficient mixing of organic and mineral soil often resulting from the activities of soil fauna; they have a relatively thick, dark, mineral Ah horizon enriched with organic matter.

Moders are humus forms with properties that transition between those of mors and mulls.  The orders can be further divided into groups described by horizon types (such as L, F, H, Ah), which indicate the stage of decomposition, or include descriptors for composition (such as woody, fungal mossy).



An example of a fungal mor humus form.
Image by C.McNalty

An example of a woody mor humus form.
Image by C.McNalty

The activities of different organisms and the interactions among them have direct consequences for carbon dynamics, as they are the agents driving rates of mineralization and respiration, as well as stabilization, and sequestration. Although the actions of soil fauna mediate the cycling and storage of carbon, the direct effects of these organisms can be difficult to study, and until now have not been included in carbon models.  In addition, most carbon models focus on predicting carbon emissions because of the great interest in greenhouse gases, and they neglect the formation and storage of soil organic matter, which is important in carbon sequestration.

An invasive earthworm and lepidoptera larva
in a boreal forest soil sample.
Image by C.McNalty

Recently, a model developed by scientists in Russia, Germany and Canada, Romul_Hum (the soil module of the individual tree forest growth model EFIMOD), integrates knowledge of humus form development, soil fauna food webs, dynamics of fungi and bacteria, and the resultant formation and stabilization of carbon in soil organic matter. Romul_Hum accounts for the many belowground interactions (fungivory, bactivory, predator-prey relationships) between soil fauna, bacteria, and fungi and how these interactions are regulated by the qualities of incoming vegetative material. Unlike other models, which treat decomposers as a homogenous group, Romul_Hum uses variations in the ratios of fungal to bacterial biomass, and the ratios of carbon to nitrogen in the fungal and bacterial biomass. Based on published soil fauna data, different types of food webs are defined for combinations of decomposers and types of horizons in humus forms. An earthworm module was developed with parameters for for food palatability, ingestion and egestion, food consumption, lifespan, excretion, and assimilation efficiency. It is especially important to understand the effects of earthworms interacting with other soil fauna activities in Canada as earthworms are invasive to Canada’s large boreal forest (see blog: Earthworm invasions in northern forests).  As earthworms spread through the Canadian boreal forest, they will change the carbon dynamics and carbon balance of the ecosystems.

Romul_Hum could be used to predict, understand, and quantify those changes. The types of changes to the forest carbon cycle from invasive earthworms is dependent on the species of earthworm(s) present. These changes can include mixing organic material in the forest floor with the mineral soil below, shifting the balance between fungal and bacterial biomass, and stabilizing carbon in soil organic matter as it moves through the gut of the earthworm, or as earthworms ingest and stabilize faeces produced by meso-fauna.  Beyond these effects, earthworms can change the food and habitat available for other groups of soil fauna, such as nematodes, mites, and springtails, affecting the survival and success of meso-faunal populations.

The Forest Floor Recovery Index
Image by D.Hoffman

By acknowledging and accounting for the complex interactions between soil faunal food webs and their habitat (humus forms),  the modelling approach of Romul_Hum provides a means to evaluate how management, and potentially climate change, affects relationships between soil fauna biodiversity and soil carbon sequestration. In Europe, humus form classification is currently being refined within the HUMUSICA project. In Canada, humus forms are used as part of a system (Forest Floor Recovery Index) to evaluate the success of reclamation after mining.

*Cindy Shaw ( is a Research Scientist, and Darrell Hoffman ( is a Forest Soil Research Assistant, for the Canadian Forest Service at Natural Resources Canada. Oleg Chertov is a scientist who has worked in Russia, Germany, and Finland, and created the Romul_Hum models with Alex Komarov


Assessing Amazonia Biodiversity: Beyond the Taxonomic Impediment

Mushrooms found on terra-firme (tropical rainforest)
on Benjamin Constant, AM, Brazil. Photo credit C.Ritter

By Camila Ritter, PhD. student, University of Gothenburg


Amazonia is the largest tropical rainforest with the highest level of species diversity in the world. However, most of what is known about patterns of biodiversity in this area is based on large-sized and well-studied organisms such as mammals, birds, amphibians, and flowering plants. Because these macro-organisms constitute just a small fraction of the world’s total biodiversity (vertebrates represent only about 0.7% of all species of eukaryotes, for instance), and no consensus has been reached on whether poorly studied taxonomic groups such as arthropods and micro-organisms follow the same distribution patterns as macro-organisms, it is urgent to put more efforts in this ‘hidden’ biodiversity.

However, to achieve this goal we must overcome a severe obstacle: the so-called taxonomic impediment. Considering that it takes, on average, 21 years from the first collection of a species until its formal description, we would have to wait another 1,200 years to catalogue all extant species. That is unacceptably slow, and we need to develop and validate new methods for faster, more cost-effective, and objective biodiversity assessments, which do not rely on manual identification of specimens. Fortunately, molecular tools have opened a new research window on biodiversity through genetic data. With methods such as metabarcoding, it is now possible to quantify phylogenetic diversity of any locality without the need for a priori classification of specimens.

In an effort to assess the main patterns of biodiversity distribution in Amazonia, we have sequenced genetic markers from both prokaryotes and eukaryotes from a range of soil and litter samples. We targeted four locations covering the different kinds of habitat (tropical rainforest, seasonal flooded forests, and naturally open areas) of the Amazonia. If we find that environmental genetic diversity and traditional taxonomic metrics are highly correlated, that would mean that biodiversity can be rapidly and cost-effectively assessed without the demand of taxonomic experts. This result would facilitate the detection and protection of areas of high biodiversity and would allow taxonomists to focus on species descriptions and the biology of the underlying organisms, rather than routine specimen identifications. If these variables, however, are found not to be correlated, it would mean that despite centuries of research we still know virtually nothing about how the great majority of the world’s biodiversity is distributed.

Naturally open area on Amazonia. These areas have an
insular distribution in “seas” of tropical forest and are
associated with white sand soil. This picture was taken on
“Reserva da Campina”, close to Manaus, AM, Brazil.
Photo credit C.Ritter

The tree shows the water mark from flood season of Várzea,
seasonally flooded forest.The mark indicates a flood height
of around 15 meters. Photo credit C.Ritter



Sunrise on the Cuieras river, “Reserva do Cuieras”,
close to Manaus, AM, Brazil. Photo credit C.Ritter

Camila Ritter taking notes from field work
Photo credit: N. Slobozian


2nd Global Soil Biodiversity Conference

Presentation screen in the Poly Grand Theatre
Nanjing International Youth Culture Centre, China.
Photo credit E.Bach

By Elizabeth Bach, Executive Director, Global Soil Biodiversity Initiative, Colorado State University


The 2nd Global Soil Biodiversity Conference (GSBC2) was held 15-18 October, 2017 in Nanjing, China.  More than 1000 scientists from 47 countries gathered at the Nanjing International Youth Culture Centre to hear about the latest soil biodiversity science, network with colleagues new and old, and share new ideas.

Applying soil biodiversity science to global policy, Dr. Ronald Vargas, the Soils and Land Management Officer at the UN Food and Agricultural Organization and Secretary of the Global Soil Partnership, and Dr. Luca Montanarella, European Commission Joint Research Centre and chair of the Intergovernmental Technical Panel on Soils, called for a formal international assessment on soil biodiversity.  There was much excitement around this call and it will be exciting to see it develop.

Conference co-chairs Prof. Ren-fang Shen (Institute of Soil Science, Chinese Academy of Science) and Prof. Yong-guan Zhu (Institute of Urban Environment, Chinese Academy of Science) shared recent developments in soil biodiversity research in China.  Soil biology research has accelerated tremendously in the past 5 years in China and support for future work is strong.  The China Soil Microbiome Initiative is a more than USD$35 Million project to systematically survey and sequence soil microbial diversity across China and link this biodiversity with key ecosystem services including nutrient cycling, crop production, and carbon storage.  Leaders in the Soil Science Institute of the Chinese Academy of Science are seeking additional funding to extend support to include soil animals as well.  In addition, the hundreds of Chinses graduate students, postdocs, lectures, and professors in attendance demonstrated deep knowledge, enthusiasm, and curiosity driving soil biodiversity research in China on topics ranging from agricultural production, ecosystem functioning, and restoration.  In addition, the 19th National Congress of the Communist Party of China happened in Beijing the same week as GSBC2, and President Xi Jinping specifically included soil and ecological concerns as priorities for the Chinese agenda.

Dr. Diana Wall, Scientific Chair for the Global Soil Biodiversity Initiative (Colorado State University), highlighted key Global Soil Biodiversity Initiative (GSBI) accomplishments since the 1st GSBC including publication of the Global Soil Biodiversity Atlas (including a new website), publication of many prominent papers, establishment and funding of working groups like sWORM, and incorporation of soil biodiversity into international policy documents including the Global Land Outlook (UNCCD), Global Assessment on Land Degradation and Restoration (Intergovernental Platform on Biodiversity & Ecosystem Services, IPBES, expected spring 2018), and Global Assessment on Soil Biodiversity (IPBES, expected spring 2018).

Zhongshan Mountain (Purple Mountain) is
east of Nanjing, China. Photo credit E.Bach

Nanjing International Youth Culture Centre
hosted GSBC2. Photo credit E.Bach

Participants enjoy a poster session at GSBC2.
Photo credit E.Bach

Xuanwu gate to the old Nanjing City Wall
Photo credit E.Bach

The GSBC2 featured research from around the world, including Europe, North America, South America (Brazil), Australia, and Africa.  The European Joint Research Centre is gearing up for the 2018 Land Use/Land Cover Frame Survey (LUCAS) which will include DNA sequencing analysis for the first time.  Dr. Brajesh Singh shared the first published results from the Biomes of Australian Soil Environments survey, showing distribution of bacteria and fungi across the nation. Brazilian scientists shared emerging work focused on soil microbes and fauna in the Amazonia region, investigating impacts of deforestation, agricultural management, and ecological restoration.  Several Canadian and American scientists highlighted how soil biodiversity interfaces with global challenges including climate change, providing ecosystem services, ecological restoration success, and plant evolution.

In addition to talks and posters, the conference featured several workshops and roundtables to share knowledge between established and early career scientists.  Writing workshops were led by Dr. Wim van der Putten (Netherlands Institute for Ecology, NIOO-KNAW), Dr. Karl Ritz (University of Nottingham), and Dr. Josh Schimel (University of California Santa Barbara).  In addition, there was an open evening social with the editors of Soil Biology & Biochemistry.  Dr. Stefan Geisen (NIOO-KNAW) organized a roundtable featuring experts sharing best practices and emerging methods for studying soil organisms including bacteria, viruses, protists, nematodes, mites, molecular approaches for mesofauna, and working across disciplines and beyond the scientific world.  Ting-wen Chen (University of Göttingen, Germany), Dr. Meixiang Gao (Harbin Normal University, China), Dr. Stefan Scheu (University of Göttingen, Germany), and Dr. Tancredi Caruso (Queen’s University of Belfast, Northern Ireland, UK) organized a roundtable focused on ecological theory and soil biota, which spurred exciting discussion late into the evening.  Dr. Nadia Soudzilovskaia (Leiden University, Netherlands) and Dr. Gerlinde de Deyn (Wageningen University, Netherlands) organized a roundtable linking soil biodiversity to ecosystem functioning and provisioning of ecosystem services.  These provided important opportunities for attendees to interact in focused small-groups and develop important career skills.


Look at the full program, including keynote abstracts here: The 3rd Global Soil Biodiversity Conference will likely happen in 2020, stay tuned for more information! Tags: 

Soil modeling for everyone: From concept to simulations

By Katherine Todd-Brown, USA @KatheMathBio

Computational and mathematical modeling are increasingly in demand as society tries to apply current scientific understanding to future soil management practices. The flux of carbon from the soil to atmosphere in particular is a major uncertainty of future climate projections. Soil carbon is also a critical component of soil health. In addition, many funding agencies are interested in coupling experimental and modeling approaches to better target scientific inquiry and extrapolate into murky futures.

Computational and mathematical models are generally thought of as tools for projecting into the future, but they can also help design and run experiments that might otherwise be costly or even impossible to carry out in the lab/field. You can turn off gravity in a model. While I say this with a bit of tongue in check, imagine directly exploring the effects of capillary action vs gravitational forces in soil wetting. Something that is completely impractical in the real world can become almost trivial in the computer.

Models are frequently seen as this magic box that, if you can just figure out how to plug your data in, will tell the rest of the world how critically important your research is. As a modeler, I’ve found that my colleagues who work primarily in the field and/or on the bench are frequently at a bit of a loss how to wave this magic wand. Fully characterizing a site to run an off-the-shelf model like DayCent can be overwhelming or just plain intractable depending on the location. Plus experimental treatments or particular measurements will often not have direct representation in these off-the-shelf models, precisely because their effects are novel or uncertain and thus of interest.


So where does that leave us?

Fortunately, if you design experiments and interoperate their results, you are most of the way there; even if you are a field or laboratory scientist who hasn’t written mathematical equation since your college math class. Numerical models are formalizations of conceptual frameworks that every scientists use on a daily basis. We tell stories of how we think the world works in our grant proposals and manuscripts. Those stories are models in their most basic forms; English, Mandarin, mathematics, and computer code are just a different languages that can be used to convey some scientific understanding to the broader world. By telling stories of how we think the world works, scientists are creating models.

The main difference between conceptual models and numerical models is precision. Human language is generally very fuzzy leaving quite a bit of room for interpretation to the listener. Mathematics, on the other hand, is extremely precise. It’s one thing to say that the change in soil carbon is the difference between the inputs and outputs, and another to state: . This mathematical formulation has several embedded assumptions in it like: soil carbon does not change with space, and inputs change over time but that the decay rate doesn’t. Mathematics does not allow you to sweep anything under the rug, instead you must explicitly state what your assumptions are and under what conditions those hold true. While you can be as precise with an English description of your conceptual framework, it can be challenging to attain the level of detail you get from a mathematical formalization. But underpinning all this is still your original conceptual understanding of the research system.


On a practical level how do you move from this conceptual framework to a precise mathematical formulation?

One way to do this is to collaborate with a mathematician or computer scientists. Be aware however that this will frequently require several conversations since you are likely talking with someone who has no soils training. The nutrient, parent material, and biological differences between a tropical oxisol and temperate molisol might be obvious to you but not to someone trained to locate bifurcation points in dynamical systems. Often times the different soil conditions will tie directly to different mathematical assumptions in ways that are not obvious at first glance. Lobbing your data over the discipline dividing wall won’t cut it. Instead, these collaborations require a lively and ongoing exchange.

Alternatively you can develop your own model. Frequently if you have a strong hypothesis driven experimental design, you can frame this hypothesis as a numerical model and probably already do on some level. Many statistical tests like linear regressions are, themselves, numerical models. A common problem with these statistical tests is that they assume normally distributed data and linear relationships, frequently not the case in soil systems. Diagramming the mass or energy path you are interested in will often lead to a set of differential equations. Once you have this mathematical formalization you can apply model-data integration techniques to fit descriptive parameters to your data.

By using numerical models informed by your scientific understanding you can dramatically increase the power of your statistical tests, compete different hypothesis, explore the relative importance of entwined mechanisms, and extrapolate findings into management scenarios. Models allow you to convey your scientific understanding, supported by your experimental data, with a precision that is difficult to achieve in a standard scientific narrative. While the end goal is scientific understanding not mathematical poetry, models are playing an increasingly important role in biogeochemistry.


Mathematics is the language of size, shape and order […] ~ Lancelot Hogben (1936)


Kathe Todd-Brown is a computational biogeochemist at the Pacific Northwest National Laboratory in Richland, Washington. She initially trained as a mathematician but it was a bit too clean. She transitioned to soil carbon cycling and couldn’t be happier to work in one of the most interesting systems on the planet.



An Experience with Interntaional Soil Reserach Networks

Collaborators discuss grassland soil carbon research
near Hulanbuir, China. Photo credit Jiqiong Zhou

By Adam Cobb, Postdotoral Researcher, Oklahoma State Univeristy, USA


There is nothing quite like touching new soil. The soils of China are ancient, but this is my first chance to grab a handful. After leaving our hotel in Beijing at 4am, flying for three hours, and driving on a bombed-out road for another hour, we are in the steppe grasslands of Inner Mongolia in Northwest China.

We are here – with chestnut-colored soil in our hands – because of Jiqiong Zhou (soon to be Dr. Zhou) and her advisor, Dr. Yingjun Zhang. Sixteen months previously, Jiqiong (who also goes by Jolie) emailed Dr. Gail Wilson and asked if she could spend nine months with us at Oklahoma State University. Jiqiong’s research links above- and belowground influences on grasslands with particular emphasis on plant-microbial relationships, such as rhizobia/meliloti and arbuscular mycorrhizal symbioses, which is precisely Gail’s area of expertise.

This PhD student exchange and research collaboration has been amazing. While Jolie was with us, she had the chance to visit the Konza Prairie Biological Research Station (NSF-LTER site) near Manhattan, Kansas, USA. We went up to establish grassland restoration plots, but took time to show her around the station, and particularly to see the bison. Later, she told us how lucky she thinks we are to research in such productive grasslands.

Back in China, her research grasslands are not as productive. With typically 300-400mm annual precipitation, they are a contrast to our tallgrass prairie systems. However, these semi-arid grasslands are critically important, not just for local sheepherders, but for soil ecologists. Climate models predict many mesic grasslands will become more arid during the 21st Century. It is crucial that we untangle the belowground drivers influencing aboveground productivity, plant species diversity, and ecosystem functions in semi-arid grasslands, if we are to manage our global grazing resources in a warmer and drier world.

Cattle grazing in the meadow steppe grasslands
of Inner Mongolia, China. Photo credit Jiqiong Zhou

Bison on Konza Prairie Biological Research Station
near Manhattan, Kansas, USA. Photo credit Adam Cobb

This is where Jolie, and her colleagues in Dr. Zhang’s lab, conduct their valuable soil research. We walk across her plots, and see the yellow-flowered alfalfa (Medicago falcata) she established to rehabilitate these overgrazed areas. Belowground, she assessed how these reseeded legumes influence nutrient cycling, soil microbial communities, and soil metabolites. Today is a hot and sunny one, and we remark at how much data she has collected in these plots, often working by herself.

Jolie’s research is the combination of mechanistic assessments (e.g., microbial genomics) and applied questions. As part of the China Agricultural University’s College of Animal Science and Technology, she is keenly aware that these grasslands are provisioning ecosystems. Grazing treatments (real or simulated) are always part of the research design. Jolie is passionate about reseeded alfalfa because it can regenerate the landscape, increase incomes, and improve soil health.

Gail and I had an amazing experience with Jolie, Dr. Zhang, and their associates. We learned more about their culture, and we touched China’s soil. We are currently planning paired experiments assessing grassland plant and soil dynamics in both China and the USA. These collaborative research networks are key as we strive to conserve and restore global soils.


Building mud castles: A perspective from brick laying termites

Termite mound construction using bricks.
Image by N. Zachariah

By Nikita Zachariah, Graduate Student, Centre for Ecological Science, Indian Institute of Science, Bengaluru, India


Walking in the wild or even in a metro city like Bengaluru you are sure to find animal homes in all their grandeur — bower bird nests, bee hives, spider webs and termite mounds. These aesthetically designed structures have always fascinated architects, naturalists and laypersons alike, yet we barely know how they are built — what are the basic building blocks or bricks in these constructions and how materials are chosen for these constructions.

In consultation with my PhD advisors Prof. Renee M. Borges and Prof. Tejas G. Murthy I decided to explore the physical, chemical and behavioural aspects of one such construction — the termite mound. Though made up of soil, termite mounds can stand in sun and rain for decades together without dissolving thanks to termite secretions that are mixed with soil during construction imparting ten fold increase to its strength. Termite mounds can house more than a million termite individuals and can reach a height of 10 metres. At a human scale this would correspond to a building 10 kilometres tall… taller than Mount Everest!! Termites construct these mounds without an architect, without a masterplan, in fact without even seeing the structure they are building. Yes, these termites are blind. Not only do termites engineer their mounds, they also engineer entire ecosystems making them drought resistant. Yet, little do we know about the basic building blocks of these mounds and what makes a geographic region conducive for mound construction.

I studied Odontotermes obesus species of termites in Bengaluru, India. It is widely distributed in the Indian subcontinent and makes mounds that are upto 2.5 meters tall. It aggregates moist soil particles into tiny balls which act as bricks during mound construction. The different castes of termites (such as major and minor termites) make different sizes of bricks which they jointly pack like golf balls in a jar with marbles filling the space between the balls thereby achieving tight packing and consequently high strength. Moreover, in the lab they were even able to use materials like glass beads for making bricks. Since termites used a totally unfamiliar material, glass beads, I was curious to know what else can they handle? In order to understand this I gave them every material I could get my hands on — metal powders, jellies, even tissue paper and paraffin wax!! To my surprise they used all the materials as long as they were able to walk and chew on them. But they do had their personal favourites, e.g. they loved granular materials over others and were equally willing to use non-familiar materials like glass beads as the familiar ones (soil). Other properties that determined the ease of handling were hydrophilic, osmotically inactive and nonhygroscopic nature, surface roughness, rigidity and presence of organic matter. These material properties along with the availability of moisture and favourable climatic conditions will determine the global geographic distribution of termites, a matter of considerable importance given their roles as ecosystem engineers. This study also takes us towards understanding how tiny termites make mounds that any engineer would envy.

Bricks made by different castes of termites.
Photo credit: N. Zachariah

Packing of large and small bricks during construction.
Photo credit: N. Zachariah


The study was published in the journal Scientific Reports ( and was featured in Science magazine (

Lab webpage: Prof. Renee M. Borges (, Prof. Tejas G. Murthy (


Biodiversity in a warmer world: lessons from soil nematodes

Predatory female nematode Clarkus sp.
Image by M. Ciobanu

By Madhav Thakur, postdoctoral researcher, German Centre for Integrative Biodiversity Research (iDIV)


John Haldane, the famous evolutionary biologist, popularized the immense diversity of beetles by writing: “The creator, if he exists, has an inordinate fondness for beetles”.  I wonder if Haldane was aware of nematodes. Nematodes are incredibly diverse and abundant tiny worms living almost everywhere on planet Earth. Some estimates point that we may have a million of nematode species on Earth. I remember Tom Bongers, a world-known nematode taxonomist, in his lectures saying every time one samples a forest soil, s/he is likely to find a new species of nematodes.

At the beginning of my PhD in 2013, I got interested in how on-going climate warming affects biodiversity. At this point, I already was familiar with the free-living nematodes in the soil, and in particular, impressed by their omnipresence in any environment.  After the consultations with my PhD supervisor, Prof. Nico Eisenhauer, I decided to investigate whether warmer soil harbours less or more diversity of nematodes. Fortunately, I was offered to investigate nematode diversity in a long-term climate warming experiment in the meadows of Cedar Creek in Minnesota, USA. This climate warming experiment was unique for two reasons: 1) Climate warming was experimentally crossed with plant diversity and, 2) Plant diversity treatments in this experiment were the part of the BigBio experiment, which is one of the oldest biodiversity experiments in the world. Further, I was very excited to work with Prof. David Tilman, who is the principal investigator of this experiment, and well-known for his contributions for our understanding of the causes and consequences of biodiversity.

Areal view of Biodiversity & Climate experiment
Image by J. Miller

Once I was able to collect nematodes and identify them with the help of colleagues (Dr. Marcel Ciobanu in particular), I started getting back at the warming and biodiversity question. Since we understand biodiversity in many different ways, I calculated many different metrics of biodiversity and see whether I could find any consistent pattern. The most striking and consistent pattern was that warming both increased and decreased nematode diversity. The key was whether the nematodes were from plant monoculture soils or from the soil of diverse plant communities. Warmer plant monocultures were lower in nematode diversity, whereas warmer diverse plant communities were higher in nematode diversity. Although these results were exciting, I did want to explore further. With the help of Dr. Oliver Purschke, I investigated whether warming also structures nematode communities in a given way. Indeed, the results revealed that nematodes were taxonomically more similar than expected in warmer soils independent of plant diversity. So even if warming increased nematode diversity in diverse plant communities, nematode communities become increasingly similar. We published these results recently in Science Advances.


The climate warming and biodiversity question will stay with ecologists for a longer time. Our results do provide some clues by using one of the most diverse and abundant organisms. At this stage, I am even more curious by the question how general are our results. Whether Haldane’s beetles, or our nematodes, biodiversity in a warmer world is very likely to be different than the past and present biodiversity.



M. P. Thakur, D. Tilman, O. Purschke, M. Ciobanu, J. Cowles, F. Isbell, P. D. Wragg, N. Eisenhauer (2017), Climate warming promotes species diversity, but with greater taxonomic redundancy, in complex environments. Science Advances 3, e1700866. Doi: 10.1126/sciadv.1700866.


Earthworm invasions in northern forests

Lumbricus terrestris
Image by E. Cameron

By Erin Cameron, Postdoctoral Researcher, Helsinki University, Finland



Finishing my paper route always took longer on rainy days when I was a kid – I had to walk slowly to avoid stepping on earthworms and occasionally I stopped to move them off the sidewalk. I would never have believed that earthworms were invasive in much of Canada and the northern United States! In those previously glaciated areas, there are no native earthworms, but instead only European earthworms that were introduced to North America with the arrival of European settlers.

Earthworm populations can only expand about 5 to 15 meters per year on their own, and consequently people play a key role in their spread. For my master’s thesis with Dr. Erin Bayne at the University of Alberta, I tried to determine how earthworms were spreading in Alberta’s boreal forest. I was sold on the research question once I realized that I’d need to canoe or kayak across lakes to test whether earthworms were most common near boat launches where anglers might dump their earthworm bait. After a couple near misses but only one capsizing incident, we found that earthworms were present at approximately 70% of the boat launches and roads sampled, but only 35% of far shores and less than 15% of forest interiors. They were also more likely to occur at older roads than more recently built roads, suggesting that earthworms were introduced by vehicle traffic (their eggs can become stuck in tire treads) not during construction of the roads.

After examining how earthworms were being introduced, we started to investigate their effects in the boreal forest. Surprisingly to most people, earthworms do not always improve soil health or benefit other organisms. When exotic earthworms invade forests where there are no native earthworms, they consume leaf litter layers, mix organic and mineral soil horizons, and affect nutrient cycling. These impacts on soil structure and ecosystem functioning can then lead to cascading effects on other organisms. In northern Alberta, we found that earthworms decreased the thickness of the leaf litter layer, reduced the abundance and diversity of microarthropods, and decreased plant biomass, depending on the species. Not all species were negatively affected though – one of the key predators of earthworms, the American robin, was more likely to occur in areas where earthworms were present.

Sampling for earthworms in the boreal forest
Image by R. Rocha

Earthworm invasions are at an earlier stage in northern boreal forests than temperate hardwood forests, where the deep burrowing and mineral soil dwelling species that cause the largest changes are more widespread. At our study sites, the most common species is a litter dwelling species called Dendrobaena octaedra. But because most people are not aware that earthworms are invasive, they continue to introduce earthworms by dumping their bait, moving soil, or not cleaning their tires when travelling to remote areas. We started a citizen science project to collect data on earthworm distributions across Alberta, which at the same time serves to increase public awareness about earthworm invasions:


However, earthworm invasions are occurring globally, rather than only in North America. We also lack data on distributions of native and exotic species of earthworms at broad scales, making it difficult to determine the key factors driving their distributions. To address this issue, we started a working group (sWORM; at the German Centre for Integrative Biodiversity Research (iDiv) to synthesize data on earthworm distributions. Let us know if you have earthworm data and want to participate!


Further reading:

Cameron EK, Bayne EM, Clapperton MJ. 2007. Human-facilitated invasion of exotic earthworms into northern boreal forests. Ecoscience 14: 482-490.

Cameron EK, Bayne EM. 2012. Invasion by a non-native ecosystem engineer alters distribution of a native predator. Diversity and Distributions 18: 1190-1198.

Craven D et al. 2016. The unseen invaders: introduced earthworms as drivers of change in plant communities in North American forests (a meta-analysis). Global Change Biology 23: 1065-1074.


The Mysterious Case of the Microbe in the Soil

Microbial biosensors sense and environmental signal and,
in response, produce a measurable reporter such as a gas.
Image by E. Fulk

By Emm Fulk, graduate student, Rice University, Houston, TX, USA


Flagella? Check. Cell wall? Check. Cheesy music? Yes, that's right. What's a microbial detective without a melodramatic theme song?

Studies of soil microbiology have traditionally - and necessarily - been conducted from the outside in. Net fluxes of nutrients and gases from the soil environment can be chemically measured. High-throughput sequencing techniques can give metagenomic data, which provides a snapshot of the general composition of a soil microbiome. This is, essentially, a stakeout - we can get a general sense of a soil  community by observing the surrounding environment and may be able to infer some activities by measuring who and what comes in and out. These strategies give us an overall picture of soil communities and their net interactions within the ecosystem but lack the spatial, temporal and chemical sensitivity to fully understand the internal  dynamics of soil microbiomes.

We need a microbe on the inside.

The idea of using living microbes as biosensors is not especially new. To survive and adapt to new environmental stresses, such as nutrient or water deprivation, microbes have evolved networks to sense these changes and adapt their metabolism accordingly. Tying these naturally-evolved systems to a measurable reporter (for example, a fluorescent protein) is a logical step for understanding how microbes interact with their environment. Think of a light bulb and a light switch. The light switch senses whether it is on or off. The light bulb reports on the ON/OFF state of the light switch. Even if you can't see the switch, you can infer whether it is on or off by looking at the light bulb. Likewise, we can detect when a particular environmental condition elicits a microbial response by monitoring the production of the reporter.

I know just the microbe for the job… he's sensitive, discreet and reports only to me.

For microbial biosensors to be useful in soil, their sensors and reporters must both be suitable for monitoring interesting  
environmental conditions - for example, drought conditions, concentrations of nitrogen or carbon species, or cell-cell  
communication signals. Sensors for -osmotic stress, nitrate, quorum sensing molecules and various heavy metals have been or are being  
developed. These systems must also be sensitive to an environmentally relevant level of the signal. If too sensitive or not sensitive  
enough, reporter production is not triggered at the right times to give useful information. Ideally, reporters need to be measurable in  
situ. Traditional fluorescent or pigmented reports can't be used because, well, you can't see them in soil. Expanding our toolbox of  
reporters would allow us to monitor cell growth as well as environmental response and to measure multiple signals.

The name's coli. Escherichia coli.

Even after engineering a useful biosensor, it remains to find suitable host microbe. Most synthetic biology is initially done in E. coli, because it is relatively well understood and easy to engineer. E. coli can also a good first organism to test biosensor function. Expanding our ability to engineer other microorganisms is a key challenge both for biosensors in soil ecology and for synthetic biology as a whole. Numerous other organisms - for example, several soil-dwelling Pseudomonas species - have successfully been engineered as biosensors. However expanding our repertoire of bacterial hosts will enable biosensors to be used in more microbial communities.

Where were the suspects on the night of the…high nitrate concentration?

Recent developments in synthetic biology have greatly expanded our ability to manipulate microbes and perform increasingly difficult computations. For example, microbes can now be programmed to produce a reporter only if both signal 1 AND signal 2 are present. Other logic functions (1 OR 2, 1 AND NOT 2, etc.) could allow for studying specific combinations of environmental conditions, such as in hot spots or during hot moments.

In standard biosensors without memory, microbes produce a reporter directly
in proportion to the level of signal. In biosensors with memory,
the reporter is induced by the first instance of the environmental cue and remains on thereafter.
The top cartoon shows how these two scenarios compare at the level of individual microbes.
The bottom graph illustrates how reporter production differs at the whole-sample level.
Image by E. Fulk

Additionally, microbes can now be programmed to "remember" certain events. By using a class of enzymes called DNA recombinases, permanent changes can be made in the DNA of a microbial biosensor in response to the environmental signal. This change catalyzed by the DNA recombinase - specifically, a reversal of a short section of DNA - can be induced by the environmental signal and results in permanent, stable, and heritable expression of the reporter. Thus, the microbes have memory of the signal. Think again about the light switch - once the light switch is turned on for the first time, the light bulb is permanently on whether the switch turns off again. This could be useful for studying long-term behavior of microbes where consistent measuring of the signal is not always possible, or for situations where suitable reporters are limited (since the recombinase-mediated reversal of the DNA sequence is a sort of reporter in itself).

It was the Rhizobia! Case closed.

Well, not quite. This is just a small slice of the synthetic biology tools that may be of interest, and there are still engineering challenges to be addressed. Most crucially, however, we need to find the right soil ecology mystery to solve.



Urban Expansion, Land Cover, and Soil Ecosystem Services

By Ciro Gardi, Scientific Officer, Animal & Plant Health Unit, European Food Safety Authority, Parma, Italy


We are aware, especially the readers of this blog, of the immeasurable value of soil and of its unique and essential role. The focus of the Global Soil Biodiversity Initiative is the variety of living forms that soil can host, and it is clear that in order to have soil biodiversity we need to have soil. In other words, we will not be able to protect soil biodiversity if we are not protecting the soil as whole. Unfortunately, there are several processes leading to soil degradation and the intensity and combination of them vary across the globe.

One of the most irreversible process, often overlooked, is represented by soil sealing, consequent the expansion of urban and industrial areas or the construction of transport infrastructures. The intensity of this process can be extremely high, especially in the countries with fast economic and/or demographic growth, and often it occurs at the expenses of the most valuable and fertile soils. After the soil is sealed, due to the construction of buildings, roads, etc., all the soil ecosystem services are lost or severely compromised. Not only to the capacity of soil to be used for agriculture, but also its capability to infiltrate water, to store carbon, etc.

A newly published book, Urban Expansion, Land Cover and Soil Ecosystem Services, is an accurate overview of the impact of urban expansion processes on the provision of soil ecosystem services. From the analysis of the magnitude and intensity of these processes at global scale, to the assessment of the impact of soil sealing and land take on the capability of soil to produce food and biomass. The role of soil in agricultural production is probably the most obvious: according to FAO, 95% of our food derive directly or indirectly from soil, and by to 2050 we will need to increase the food production by 70%. There are however other soil ecosystem services essential for human wellbeing, and more in general for the protection of life on the Earth. The regulation of water cycle and most terrestrial biogeochemical cycles rely on soil. The existences of the majority of terrestrial ecosystems also depend on soil.

It is essential then to have a more responsible use and planning of this strategic and pivotal non-renewable resource, sharing our knowledge of its values and role, with policy makers, land use and urban planners, but also citizens. In Europe, a public campaign has been recently launched (People4Soil - ) to request the declaration of a soil protection Directive at the EU level. This is an example of the efforts of awareness raising that made possible to spread among citizens the comprehension of the role of soil, as essential element of the natural capital.

Often new residential, commercial
or industrial buildings are realised
only for speculative reasons and remain
unused for ever. Photo credit: Ciro Gardi

Soil sealing prevents, or strongly
compromise all soil ecosystem functions.
This is an example of an exception:
the vigour of this plant is able to break
the asphalt coverage sealing the soil.
Photo credit: Ciro Gardi

An example of a construction site.
Soil is subject to several types of
degradation processes: compaction,
contamination and finally sealing.
Photo credit: Ciro Gardi



Learn more about the book here:


Cast in clay

By Max Helmberger, graduate student, New York State Agricultural Experiment Station, Geneva, New York, USA


Growing up an only child on a dirt road in the Northern Minnesota woods, I spent a lot of time outside, overturning the many thousands of glacier-strewn rocks around my house (at least the ones small enough to move) and gazing in awe at the centipedes, isopods, and invasive European earthworms underneath (which were promptly relocated to our compost bin). My first interest in soil in an academic context came the year after I graduated from high school, when I took a soil science course at my local community college. That class instilled in me a firm conviction that soil is humankind's most important natural resource, and clued me in to the fact that, in soil, there's far more than meets the eye.

After transferring to the University of Minnesota in Duluth to major in Biology, I took an entomology class and worked as a research assistant in an aboveground plant-insect ecology lab. I had always loved insects, arthropods, and invertebrates in general, and I greatly enjoyed the coursework and research experience, but wanted to connect it with my love of soil. I started searching Google Scholar for “soil arthropods”, and a few dozen soil ecology articles later, I was hooked, and I eventually sought and out and accepted a M.S. position in the lab of Dr. Kyle Wickings at the New York State Agricultural Experiment Station, a satellite campus of Cornell University located in Geneva, New York. What I didn't know as I was reading all those papers, was that I would end up reporting all the knowledge I gained from them in a most unusual way.

Clay animation, for those unaware, is a form of stop-motion animation in which clay models are photographed, moved slightly, photographed again, and so on. The pictures are strung together and played in rapid succession to give the appearance of movement. Wallace and Gromit is arguably the most famous example of the medium. When I was 7 years old, my grandmother took me to an hour-long class in clay animation. I made a video of an anthropomorphic flower dancing to some sort of classical music riff. The VHS tape is certainly hiding somewhere in my house. Fast forward to my undergraduate entomology class, and I drew on those old memories for the course's final project to make a clay animation video on the life cycle of the gallmaking fly Eurosta solidaginis, the main study organism of my research advisors’ laboratory. The video was crude by my current standards, but got me a good grade in the class nevertheless.

At Cornell, where I'm currently working on my M.S. in Entomology, I’ve had the opportunity to draw from a unique funding source called the Extension/Outreach assistantship. Instead of working off a grant or being a teaching assistant (something difficult to do when based at a satellite research campus rather than the main university campus), I earned my stipend via progress on a variety of extension and outreach projects of my advisor's and my own devising. When applying for the assistantship and listing my project objectives, my advisor and some of the faculty members on the Extension/Outreach assistantship committee were skeptical until they saw the E. solidaginis video as a proof-of-concept, and in Spring semester of 2017, I was off to the races. To avoid some of the mistakes I made with my previous clay animation, I was very careful with how I went about planning and "filming" the videos. I wrote out all of my narration in advance as well a clear script of what specific actions I would portray. Then, I timed myself reciting the narration for each scene, and so could know in advance how many frames of animation I needed. This, combined with a nice DSLR camera as a Hanukkah present from my parents, would allow for much more cohesive and polished videos than my tale of the gallmaking fly. In the end, I produced three videos, Life Cycle of Entomopathogenic Nematodes, The Soil Food Web, and Ecosystem Services in Agriculture (though the latter includes some functions performed by aboveground organisms). My funding next semester will be from the same Extension/Outreach assistantship, so I plan to produce at least one more video in addition to my other projects, possibly two. From start to finish (writing the script, creating the models, creating the set, taking the photographs, editing the video, and recording the narration), each video took me between 15 and 20 hours to make. Each video consists of 350-400 individual images, with some being repeated here and there. They’ve been received well in the department, and the entomopathogenic nematode video has even been incorporated into several extension talks. The rest of the videos have ben showcased in a few classroom settings, and I am hoping to further expand their reach. I plan to make at least two additional videos in the fall, as I will again be funded through an Extension/Outreach assistantship.

My ultimate goal is that these videos provide an accessible way of communicating soil ecology and biodiversity to lay audiences, especially young ones. Despite being the prototypical "science nerd" growing up, and being an avid consumer of books, documentaries, and Web resources about the natural world, many of the soil animals I read about during my first forays into the primary literature were completely unknown to me. I had no idea there were mites beyond dust mites and the various parasitic taxa, and certainly didn't know there were any mites as cute as a galumnid oribatid. I had never even heard of diplurans, symphylans, pauropods, and some of the other more obscure soil organisms. I knew what a pseudoscorpion was, but didn’t know I could find them in the peat bog less than a mile from my house. And that rubbed me the wrong way. It's hard for young people to learn about the marvels of soil biodiversity, especially on their own. I know that 7 year-old Max would have gotten much more out of these videos than from one about a dancing flower, and he would have started playing around with Tullgren funnels much earlier than junior year of college. As such, if you enjoy my videos, I encourage you to share them however and wherever you like. They are available online as a YouTube playlist.


Digging deeper in urban ecology: the urban ecosystem convergence hypothesis

By Dietrich Epp Schmidt, Graduate Reserach Assistant, University of Maryland

“Even the mightiest of us return to dust, they say. Nothing remains but these shattered fragments of their kingdom… But that's not really the point, is it? These shattered fragments remain- that's the point. We look upon the magnificent temples and stelae and ball courts of Caracol in awe. There's no despair here. The Maya built something astounding and permanent. Look on our works, ye mighty, and revere. The ancient Maya speak to the twenty-first century through those temples and say: We did something amazing here.

What will our descendants think when they come upon Chalillo [dam]? When they scrape away the deep layer of dirt covering its stepping-stone facade, what will they make of the dogleg design, the Chinese gauges, the long-stopped turbines? What will they make of the skeletons and fossils of birds long gone? Will they connect the two?"


-Bruce Barcott, The last flight of the Scarlet Macaw


For a very long time, humans have experienced the world as having two fundamental domains: that which we understand as being under human control, and that which is not. Canonically, we call the latter “natural” and the former “unnatural;” and while the broader society may perceive there to be a distinction between the two, we know that our existence has an impact on ecological process, and vice versa. Our social processes, which are as basic to us as defining our identity and competing for status within our community, are intricately linked to our individual and global consumption of resources. Our economic trade is itself an ecological process that transports resources and organisms across the globe [see telecoupling]; our civilization alters community composition and function wherever we can make a living, all the while affecting global biogeochemical cycles. Our framework for understanding the role of social process on ecosystem function is just in its infancy, and there is no vernacular language for describing the built environment as an ecosystem. This is not how our society understands the environments we inhabit. One goal of the Global Urban Soil Ecology and Education Network (GLUSEEN) is to increase the exposure of urban citizens to the important role that soils play in maintaining ecosystem health.

Within the discipline of ecology, there exist frameworks to describe the outcomes of human behavior in terms of ecosystem process. For instance, biotic homogenization (BH) describes a process of convergence among biotic communities; generally communities become more similar (converge) when endemic specialist species are extirpated and generalist species come to dominate. Convergence is a process that is often applied to understanding the effect of both urbanization and agriculture, where the implied (or assumed) mechanisms are generally anthropogenic disturbance and/or facilitated dispersal. In this instance, BH describes how land-use conversion (habitat loss) drives local extinctions, while the cultivation of exotic species facilitates the dispersal of a common set of organisms. In this context, BH helps to explain the paradox of high urban and peri-urban biodiversity concomitant with significant global biodiversity loss. Endemic species go extinct, while opportunists thrive in the human-disturbed landscape (see, for example, this). To bring the mechanisms into focus, BH has been reformulated somewhat as the Urban Ecosystem Convergence Hypothesis, which relates structural changes in the built environment to changes in community process. It predicts that if urban landscapes are constructed and maintained in a similar manner (causing a convergence of habitat characteristics across biomes), then their biotic community and ecosystem processes should converge as well. For example, in their paper entitled “Ecological homogenization of urban USA,” Groffman et al. show that the practice of maintaining irrigated lawns causes a convergence of biophysical conditions across biomes within urban areas in the United States; in temperate forest systems, land-use conversion to lawns increased surface temperatures by reducing shade and evaporative cooling that normally occurs in the canopy. Whereas irrigating arid land for lawns increased evaporative cooling at the ground level, causing the two environments to converge with respect to temperature as well as humidity.

Image recreated from Pouyat et al., 2017
Journal of Urban Ecology.

The Global Urban Soil Ecology and Education Network (GLUSEEN) applied the Urban Ecosystem Convergence Hypothesis to urban soils. We sampled from soils that occurred within four different land-uses, which were categorized to represent land-use histories that are typical of urbanized landscapes (published here). These four categories were reference, remnant, turf and ruderal. Reference sites served as our control; they were sites located outside of the urban matrix, which are representative of the historic state of the ecosystem and are being managed to mitigate human impact. Many reference sites were areas set aside for habitat conservation. Remnant sites are similar to reference sites in community structure but occur within the urban matrix, and thus exposed to urban environmental factors. Turf sites were defined as sites under management to maintain a turf-grass system, which include municipal, residential, or park lawns. Ruderal sites were defined as sites that have experienced recent and substantial disruption of the soil profile, and typically were areas with a history of demolition or construction activity. Using these land-use and cover categorizations, we asked whether specific types of land-use and cover (both largely an outcome of cultural processes) caused physicochemical properties and microbial communities of soils to converge; and whether these changes result in a convergence of function among these soils.

Within-group variance of edaphic factors, among land-use;
2a shows the convergence of soil pH, OC, and N
under turf and ruderal land-use relative to the reference;
2b shows the divergence of P and K under turf
and ruderal land-use relative to the reference.
Recreated from: Pouyat et al., 2015.

As an assessment of the soil habitat characteristics, and to test the first question, we measured edaphic features such as phosphorus (P), nitrogen (N), and potassium (K) availability, as well as other characteristics such as organic carbon (OC) and pH. To quantify and identify the archaeal, bacterial, and fungal community, and to test the second question, we used quantitative PCR and amplicon sequencing. And finally, as a test of soil function, we conducted a decomposition experiment using tea bags in each of the study sites (see here). First, we found that in fact some physicochemical properties of soils converged under turf and ruderal land-use and cover types. Soil pH, OC, and N in particular converged. However, not all characteristics converged as K and P actually diverged under urban land-use (Figure 1).  We believe that it’s likely that cultural differences in how fertilizers are formulated (N vs N:P vs N:P:K fertilizers) and the variability in their rates of application may explain the increased variability among P and K nutrients; while N is also enriched systematically by fossil fuel combustion that leads to consistent atmospheric deposition of N in urban areas (and thus convergence). The convergence of soil pH is likely related to the widespread use of concrete, and the resulting concrete dust in urban areas; the calcium oxides and carbonates found in concrete effectively act as a liming agent as they dissolve, buffering soil pH towards a more alkaline condition. And finally, while specific land-use and cover types might have differing effects on the soil OC concentration, the effects within each land-use are consistent; disturbances often result in lower OC, while irrigation and fertilization in the absence of disturbance may actually increase carbon storage in soils. Thus, cultural factors may drive convergence of some habitat characteristics while causing other habitat characteristics to diverge.

Within-group variance of the fungal, bacterial, and archaeal
communities; the fungi converge in ruderal sites
relative to reference,the bacteria do not converge,
and the archaea converge in the turf and ruderal sites.
Recreated from Epp Schmidt et al., 2017.

Our next question was whether communities of organisms living in the soil converge under similar land-use and cover. We found that of the three phylogenetic domains making up the soil microbial community, the archaea and fungi exhibited a marked convergence, while bacteria did not (published here). We also showed that this convergence may be driven by different ecological factors. For example, we found that convergence in the fungal community was largely due to the loss of ectomycorrhizal fungi (ECM), while the convergence of archaeal communities was due to the increased abundance of ammonia oxidizing archaea. ECM function as symbionts with woody plants, and thus are highly reliant on the abundance of their host species. When land is converted from forest to any non-forested urban land-use, it appears that there is no longer viable habitat for most of these species. Archaea, on the other hand, actually increased in overall abundance under lawn use, and their community became dominated by organisms that derive energy from the oxidation of reduced nitrogen species. This means that, unlike for fungi, the N enrichment of urban environments is favorable to these members of the archaeal community. Moreover, since absolute abundance increased and richness also increased, it appears to be the case that the metabolic differentiation among archaea allowed convergence to occur without competitive exclusion. Therefore our dataset demonstrates the two mechanisms by which convergence might happen; a loss of unique species, or an increased dominance of just a few species that can be found in all sites. It is of course possible that both mechanisms operate in tandem to cause convergence.

The urban landscape represents the zenith of human development; it is the cultural hub of our civilization and the control center for our social process. It is also the area with the highest land-use intensity, and therefore is the most significantly disrupted ecosystem. With our dataset, we are able to show some effects that human culture has on ecosystems; the cultural process that drives the globalization of our economy and the homogenization of our global culture also has global impacts on ecosystem process via the local decisions that land managers make. Our data demonstrates that human culture may cause either a convergence (soil pH, N and OC) or divergence (P and K) of soil habitat characteristics, and that microbial communities may (fungi, and archaeal) or may not (bacteria) converge as a result. We were also able to determine multiple mechanisms that drive convergence. The fungi likely converged due to our impacts on cover (reducing the abundance of host species), while the archaeal likely converged because of our N enrichment of the urban landscape (enhancing the fitness of organisms that rely on certain forms of N metabolism). Thus there are specific interactions between human alteration of the landscape and biotic community response. The impacts of urbanization on the function and makeup of the biotic community may be long lasting, and indeed, might even outlive civilization itself. In 2008, when he published his book The last flight of the scarlet macaw, Bruce Barcott could scarcely have known that within a decade scientists studying the Mesoamerican landscape would be using the color of tree leaves (by reflecting laser light off of them from high altitudes) to discover the location of lost ancient Mayan structures. It is remarkable that urban centers, constructed and abandoned nearly a millennium ago, can still be discovered using their legacy impact on the biotic community.


Read the full manuscript here:

Epp Schmidt, D. et al. 2017. Urbanization erodes ectomycorrhizal fungal diversity and may cause microbial communities to converge. Nature Ecology & Evolution doi:10.1038/s41559-017-0123


Soil ecologists define research priorities

By Nico Eisenhauer, Professor for Experimental Interaction Ecology, German Centre for Integrative Biodiversity Research

Many, if not most, of the ecosystems on Earth are dependent on, or substantially influenced by, interactions and processes occurring within and among the planet’s soils. The remarkable biodiversity harbored in soil provides essential ecosystem services, and the sustainable management of soils has attracted ever-increasing scientific attention. Although soil ecology emerged as an independent field of research many decades ago, and we have gained important insights into the functioning of soils, there still are fundamental aspects that need to be better understood to ensure that the ecosystem services that soils provide are not lost and that soils can be used in a sustainable way. In a recent Opinion Paper (Eisenhauer et al. 2017;, we highlight some of the major knowledge gaps that should be prioritized in soil ecological research. These research priorities were compiled based on an online survey of 32 editors of Pedobiologia – Journal of Soil Ecology. The questions were categorized into four themes:

(1) soil biodiversity and biogeography;

(2) interactions and the functioning of ecosystems;

(3) global change and soil management;

(4) new directions.

While some of the identified barriers to progress were technological in nature, many respondents cited a need for substantial leadership and goodwill among members of the soil ecology research community, including the need for multi-institutional partnerships, and had substantial concerns regarding the loss of taxonomic expertise. Global efforts such as the Global Soil Biodiversity Initiative suggest that meaningful collaborative endeavors among researchers could be possible and may represent a starting point from which to build this concerted effort to address the questions presented in our Opinion Paper.

Interaction in the soil caught in the act: a predatory mite from the family Bdellidae
feeding on the springtail Sminthurinus elegans. Image by Andy Murray.


Eisenhauer N, Antunes PM, Bennett AE, Birkhofer K, Bissett A, Bowker MA, Caruso T, Chen B, Coleman DC, de Boer W, de Ruiter P, DeLuca TH, Frati F, Griffiths BS, Hart MM, Hättenschwiler S, Haimi J, Heethoff M, Kaneko N, Kelly LC, Leinaas HP, Lindo Z, Macdonald C, Rillig MC, Ruess L, Scheu S, Schmidt O, Seastedt TR, van Straalen NM, Tiunov AV, Zimmer M, Powell JR (2017) Priorities for research in soil ecology. Pedobiologia 63: 1-7.