The Kaspari Lab

As ecosystems heat and green, ant abundance and diversity increases; but too much heat and these communities lose colonies and species.

One paradox in the recent flurry of papers reporting insect declines is that insects—ectotherms that rely on external sources of heat—are often predicted to benefit as their environment warms. In an open access paper accepted as a Report in the journal Ecology  our team of ecologists—including Michael Weiser, Jelena Bujan, Karl Roeder, and Kirsten deBeurs—all from the University of Oklahoma, help resolve that paradox.

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Ants are diverse and abundant in terrestrial ecosystems. They act as nature’s cleanup squad, and are threaded throughout her food webs. Understanding how their abundance and diversity is changing as Earth heats up is a major challenge to ecologists. Images used by permission of Alex Wild @Myrmecos.

In a resurvey of 34 North American ecosystems after 20 years—both surveys funded by the National Science Foundation and that contribute to the NEON continental observatory—ant communities from deserts to forest have increased their abundance and diversity, but only up to a point.


The ants living in 34 ecosystems across North America were first surveyed in the 1990s, and then again 20 years later.


The desert, tundra, grassland and forests resampled by the Kaspari lab as we explored 20-year changes in the ant communities. Image by Karl Roeder.

To get these ecological “before” and “after” pictures, our team of post docs, graduate students, and undergraduates, traveled across North America in 2016 and 2017. We used the same methods of sampling ants from the same field stations, parks, and forest service districts that Weiser and I, along with Leanne and Alfonso Alonso, had sampled back in the mid 90s.

Temperature and NAP change

The 34 sites were on average 1°C warmer with higher estimates of plant production. Both trends are predicted to increase the numbers of ant colonies, and numbers of ant species in the ecosystem.

Twenty years later, these sites were different in important ways. First, they averaged 1°C warmer, with some much warmer than that. This evidence of global heating, differentially expressed, allowed us to evaluate a key theory underlying global change. that systems where ecotherms run the show should increase their activity and production—in fact accelerate it—before they suddenly crash. Second, they tended to be more productive–that is plants were producing more food, in the form of sugars, than before.


A graphical prediction from Thermal Performance Theory, that systems accelerate before they crash. In this case, it predicts that smaller increases in temperature are likely to enhance ant abundance, but bigger temperature increases will cause abundance to suddenly decline.

This second snapshot of the ant communities revealed that ant abundance at most sites had increased, and increases in the number of ant species had increased even more consistently.

Community Change

The distribution of observed 20-year changes in North American ant communities. Left of the blue line, communities are losing colonies and species. To the right, they are gaining.

Ant colonies tended to be more abundant in ecosystems that had warmed by a degree or so. However, those few that had warmed more—and some forest sites were up to 2.5°C hotter—suffered big declines in colony numbers. This result supports an important ecological theory: that when you warm a system composed of ectotherms it often accelerates. But beyond a certain temperature, it crashes.


Dr. Jelena Bujan sampling a one m2 plot for ant colonies at the Jornada field station in southern New Mexico. Photograph by Josh Kouri.

The number of species found in each community—a measure of its biodiversity—was in turn sensitive to these changes in abundance. The communities that increased their abundance were able to support more biodiversity, on average 2 more species on our sample transects after 20 years.


Dr. Michael Kaspari using an aspirator, or “pooter” to suck up ants at the Ordway Swisher field station in northern Florida. Photograph by Deborah Kaspari.

Such data on insect changes—collected all in the same way and over a large geographic area—are still relatively rare, making generalizing about insect declines difficult, and contentious.

But ants are an important test case. They are ecologically dominant and widespread. Their apparent increases—when studies of say, butterflies often show decreases–also likely arise because ants tolerate higher temperatures than other insect groups. Moreover, ants tend to be diet generalists, eating plants, animals, and carrion, and many colonies can seek respite underground from the hottest parts of the day. All of these traits combine to make them relatively resistant to moderate increases in temperature.

However, the precipitous decline of ant communities at temperature increases > 1.5° C is a cause for concern. We will all be watching how this unfolds over the next 10 years.


Dr. Michael Weiser, inspects the ants active at night from a plot at the Sevilleta National Wildlife Refuge. Photo by Josh Kouri.


Arts and Sciences

Caption: Debby Kaspari (of Drawing the Motmot) provided this sketch for a seminar of mine in the 90’s, an update of the parable of the blind men and the elephant. 

One of my favorite classes is “Principles of Ecology”, a venerably old course at the University of Oklahoma whose structure and focus has long been shaped by the professor in charge that semester.

One of my tweaks to PE in Fall 2017 was the addition of a list of the Ten Principles of Ecology around which to structure the course. This experiment generally paid off, I think, as it gave students ten solid memes to take away at the end of the semester. And there was a bonus: the ten were also a great way to structure PE’s final capstone project. Students have one of three options: 1) write a 500 word letter to the editor, 2) record a 15 minute teaching video aimed at middle school students, or 3) create a work of art. Each should celebrate and makes concrete why knowledge of one of those principles is a good thing.

To my delight, an increasing number of students have been creating art. Moreover, “Option 3” consistently yields some of the most intense and novel work. I’d like to share with you and honor five of those works of art here. The following gallery could easily be three times as long.

The instructions

Create a piece of art that captures one of the ten ecological principles. Create a description of the piece that would hang next to it and that includes the name of the piece, the name of the artist, the date it was created, and a short description of the materials, the process by which you created it, and the principle it was meant to capture.

A Cog in the Moo-chine by Miranda Hannon


“The cow itself was laser etched on a ½ in thick piece of plywood, using a table saw to get the plywood the correct size. The gears were 3D printed out of PLA on a MakerBot Replicator 5th Generation. The wood was lightly stained and the cow was covered in painter’s tape to give contrast. It represents Ecology Principle #3: Organisms are chemical machines that run on energy. While the interior mechanisms may differ, we are all bound by the laws of physics. Most energy ultimately originates from the sun (Principle #2), and organisms are the machines that must process that energy into a format that’s usable for them.”


 Freevector. Cow Silhouette Graphics [Online Vector]. Retrieved December 4, 2017 from

 Pleppik. (2012) Sam’s Gears [3D Model]. Retrieved December 4, 2017 from

Organized Chaos by Makenna Hukill


“This piece represents Principle 8 of the 10 Principles of Ecology that states, “Ecosystems are organized into webs of interactions.” I never realized how important species interactions truly were until taking Principles of Ecology. Measuring and understanding species interactions is one way to determine the health of a biome or ecosystem. Also, the more species interactions there are the more stable an ecosystem is. I used String Art to illustrate species interactions. I would also like to point out that I used slices of wood in this piece. I did this purposely and with meaning. Examining and measuring details in wood slices is a way to determine if a terrestrial biome has been healthy over an extended period of time. Although this piece focuses on the importance of species interactions, the take home message is having healthy biomes and ecosystems through the examination of fine details and organized chaos.”


  • Wood (from the tree cutters at McKinley Elementary School)
  • Cotton Yarn
  • Decorative Nails or Tacks
  • “T” Brackets and Screws

Interactions by Nicole Nguyen


“Acrylic paint was used to first paint the different organisms. The different organisms were then labeled with their name painted on the backs of them. Interaction arrows were also painted with acrylic paint, and the backs of those were also labeled with the type of interaction being shown. All of this was then connected by ribbon. There were three ribbons total, which were then all tied to a rod. Another piece of ribbon was tied to the rod so that this piece of artwork could be hung up. The principle it was meant to capture was principle 7, which states that organisms interact – do things to each other – in ways that influence their abundance.”

The Sunflowers of Life by Bahar Iranpour


“Materials: As an artist, my favorite media to use is paint, specifically oil paints and watercolor. However, oil paintings take up to a week to dry, so I chose to make my project using water colors. I actually prefer using water colors when drawing nature sceneries because I like how light it appears and how easy it is to mix the colors together. It makes the painting look more realistic because nature and our ecosystem is just as mixed in together and intertwined. As you can see in the painting, there is no specific line or division between the flowers and the field, the mountains and the sky. It was my way of showing through the material I chose that everything is complete when together as a whole.

Principle: I chose principle 2– The sun is the ultimate source of energy for most ecosystems. Life runs on carbon rich sugars produced by photosynthesis; every ecosystem’s sugar output depends on how much solar energy and precipitation it receives. I thought this was the most important principle because it serves as the base for all the other principles. If there was no sun, there would essentially be no life, so we would not have any organisms, hierarchies, species interactions, etc that the rest of the principles discuss. The sun allows photosynthesis to occur in plants. These plants are then able to make glucose and grow, which provides food for herbivores. These herbivores eat the plants, grow, and then carnivores come and eat the herbivores. This cycle consistently continues, providing energy to all levels on life.

Process: I definitely wanted to paint a nature scene for the principle I chose. However, I didn’t want to just paint a sun in the middle of my art. I wanted it to be more interpretive. I chose to draw sunflowers because it reminds me of when my mom used a sunflower as an example to teach me about photosynthesis when I was a child. My grandfather always had sunflowers in his garden. I remember my mom pointing out how the sunflowers would position themselves toward the sun in order to obtain its energy and go through photosynthesis. Then at night, the sunflowers would turn around again. This process was very beautiful to me, so I wanted to depict it in my painting. That’s why I drew the sunflowers facing different directions.

This taught me to appreciate the world around me, and how something as simple as the sun does so much for us. I feel that society is so invested in other things, that we take our environment, our sun, etc for granted. One day it may go away, and all life will be gone too. This is why I chose to not make the sun so apparent and clear. I painted a faint, yellow shadow in the middle of the sky, behind the mountains and clouds, to show that the sun is present and providing energy to the plants and flowers in the painting, yet it may go away one day and not be able to provide its energy.

I drew a variety of things like flowers, grasses, plains, shrubs, mountains, water, etc to show that the sun is important to all of these and many more. The small little plants on the mountains and even the small little plants in the water all need the sun’s energy to live. I included a body of water in my painting to also show the importance of water in photosynthesis—water is also needed for photosynthesis to occur. I also wanted the painting to be mostly yellow in order to emphasize the sun and its color, which is why I chose to paint a sunflower field.”

Human. Expansion. Landscape. Permutation. by Sierra Smith


“This piece was intended to capture Principle Nine of the Ten Principles of Ecology. In short, the ninth principle states that humans have an enormous influence on the Earth’s biosphere due to the large population size and technological advances the species has achieved. For hundreds of years, this influence has been used to reshape the Earth’s landscape, scatter species, and destroy diversity. This piece depicts a side by side view of what the natural Earth looks like in comparison to an Earth shaped by humans. To create my piece, I began with a blank, white poster board and used water based, twin tip Fineline Markers. I used these markers to draw the pine trees in addition to the bison and deer herds. Also, these markers were used to outline everything from the roads to the buildings to the mountains. To color everything in I used regular Crayola colored pencils.

I began the process by using the markers to draw the outlines of the buildings and the mountains to establish the two different sides of the piece. Next, I drew the road dividing the sides, and outlined it with the gray marker. Then, I drew all of the trees surrounding the mountains. I made sure to leave a space for the deer and bison herds. There was a more open space left for the bison herd in order to create a prairie environment. I moved on to the city side of the piece by drawing windows on the buildings and coloring them in. The BP building is a representation of a time humans had a detrimental effect on the environment due to the BP oil spill. After this, I drew and colored in the river. The river that runs through the city is a darker shade than the side that runs through the natural environment because of the pollutants and junk that humans release into the water systems, causing murky water. I drew the cargo ship as a representation of one way humans have used the rivers for our advantage. Then, I created the three billboards on the sides of the road. One billboard reads: “IPhone 10 On Sale Now” to signify the advanced technology the human race has developed over time. The next reads: “Land for Sale: 20 acres” which represents the fact that humans will commonly come into a natural environment and change it to their liking, regardless of its effect on the natural life inhabiting the land. The last sign reads: “Eat low fat bison burgers” to depict the human consumption of many animals, including bison. These three signs are representations of common human-imposed environmental change described in Principle 9.”

Toward a MacroEcology of Some of the Little Things that Run the World

Caption: Five Geographical Ecologists from OU’s Dept. of Biology and investigators on our new macrosystems grant.  L-R: Michael Kaspari, Matt Miller, Michael Weiser, Katie Marshall, and Cam Siler. 

The backstory behind our latest NSF-Macrosystems funded project

Insects are among the most abundant and ecologically important animals in the biosphere. Insects pollinate plants and decompose them back into soil. They include serious crop pests and invasive species that cause countless millions in damage. A key goal of macroecology is to understand how and when the number and activity of insects change as one moves from place to place across the U.S., and why those numbers fluctuate from year to year. Such an understanding can help predict insect pest outbreaks, the spread of invasive species, and changes in an ecosystems ability to provide food and fiber and conserve soil nutrients.

Yet macroecological datasets vastly underrepresent the terrestrial invertebrates. This reflects a lack of sampling effort (boots on the ground collecting insects) and identification expertise (eyes in the lab counting and recording them).

Thanks to the NEON pitfall network, 47 invert sampling arrays now span the U.S.’s major ecosystems, collecting insects in pitfall traps. These traps–sunk flush with the soil surface–capture biweekly samples throughout the growing season of each site. We believe these samples are the first step toward a new flourishing invertebrate macroecology. But, beside the carabid beetles, these samples currently consist of jars of bugs in alcohol. For the first time, we have the potential to understand both seasonal and annual invertebrate dynamics at a continental scale. If we could just count, size, and identify all those bugs in all those jars.


The geography of the NEON pitfall trap network. (A) Distribution of monitoring sites in the NEON Pitfall Network (Note: Sites outside of continental US not shown above). (B) Example of site-specific pitfall array. (C) Illustration of individual, pooled jar of macroinvertebrates collected in pitfall array.


The second step

We call the project NEONinverts*. We will endeavor to develop two complementary technologies to turn NEON’s jars of bugs into some classic macrecological variables–abundance, diversity, and body size–and then to test some big continental questions (see below). Each NEON jar is a two-week sample from one of ten arrays from 1 of 47 sites. Only the carabid beetles have been removed.  This is an astounding library of ecological information.

In NEONinverts, we will develop two pipelines–a g’mish of technology, databasing, and best practices–that will count, size, and identify taxa from these samples. The first half of our project will be to test, hone, and validate these protocols.


The big picture of our pipeline development.

Environmental Barcoding will extract insect DNA from the alcohol in which the insects steep and use that DNA to put names on the inverts found therein. Environmental barcoding (EB) is a set of tools that allows the identification of biota, not from their tissue, but from the media in which they exist. It has been used to screen for aquatic invasive and endangered species from water samples. More recently, several studies have used EB from arthropod “biodiversity soup”—homogenized samples from insect traps.

We propose to take on the hard problem of non-destructive quantification of diversity and composition from pitfall samples using the ethanol supernatant of preserved macro-invertebrate samples. In doing so, we hope to develop a robust pipeline for characterizing taxonomic diversity not only from pitfall traps, but from any fluid preserved mixed-invertebrate sample. This includes quantifying the nature and magnitude of biases, both taxonomic and environmental.

Image Analysis is a second, complementary method for quantifying pitfall traps samples. It starts with spreading trap contents on a white gridded surface with trap labels and ruler placed in a standard location. Images are captured with an articulating system of 10 PowerShot G7 X Mark II cameras, each with a resolution of 20.1 megapixels from fixed tripod arms with constant lighting conditions. Our first goal will be to use these images to quantify abundance and body size distributions.


The low down on Image Analysis as a tool to identify bugs.

We will then use machine learning–automated algorithms for pattern recognition and classification–to explore our ability to identify taxonomic groups. While machine learning methods for classification are not as accurate as expert examination, they classify orders of magnitude faster than an expert. Like the EB methodology, one of our chief goals is to see how far we can get with current technologies. Just quantifying the the dynamics of different invertebrate orders in time and space will be a huge step forward, but we are confident we can go further than that.

Together, these two pipelines hold the promise of automate and streamlining NEON’s monitoring network, providing the first such nationwide dataset on abundance, activity, and diversity of the U.S.’s soil insects.

Answering macroecological questions with invertebrate data

These data will be potentially valuable to a variety of stakeholders: ecologists testing and refining models that predict future insect communities; land managers who want to know the likelihood of a pest eruption; conservation biologists and urban planners hoping to anticipate spread of invasive ants and beetles.

We are focusing on one uber-question: how do Earth’s great abiotic drivers–temperature, precipitation, and biogeochemistry–govern how ecological communities of individuals and species vary from place to place and over time?  The few existing arthropod datasets suggest that as one travels from deserts to rainforests, terrestrial arthropod communities vary by orders of magnitude in abundance (the number of individuals), size (mass per individual), activity (the rate at which individuals do work on the system), and diversity (the number of species/forms). A better understanding of the drivers of each–applied to groups as different as spiders, ants, collembola, and tiger beetles–should help us understand how these taxa regulate ecosystem processes like decomposition, herbivory, and seed dispersal.

Stay tuned.

*Yeah, we know. We are not completely happy with “NEONinverts”. But at least the twitter handle was available.

The Ten Principles of Ecology

I have been teaching a course called “Principles of Ecology” at OU since 1996. It was a traditional two one-hour lecture, one three-hour lab for most of those years. In 2013, I decided to flip the course, converting the lectures to workshops, and asking students to do more reading outside of class. The goal is to allow for more hands on “learning by doing” activities in the workshops, and to more tightly linked lab and fieldwork to workshop data analysis and interpretation. All in all it has been an exciting, often unnerving, but very satisfying transformation. One that is ongoing.

All along it occurred to me that the title of our course strongly implies that there is a finite number of useful principles that our students should internalize. Moreover, if the list and the principles themselves are sufficiently pithy, we should be able to cover them at the beginning of the course, rather than unveil them, one after the other, as the course proceeds. The advantage there would be that students get the big picture early, allowing us to revisit and recombine different suites of principles to build and explore new concepts and ideas. That’s the idea, at least.

I had two inspirations for this venture. One was Eugene Odum’s classic “Fundamentals of Ecology”, the famous “yellow book” that was the go-to text for much of ecology’s early years. Odum organized the book around chapters with titles that begin “Principles and concepts pertaining to….” (e.g., “Limiting factors).  He would then carve each chapter into a series of expositions each with a “Statement”, followed by an “Explanation” followed by “Examples”. I just love Odum’s book and this organization because it fits so well how I organize my own thoughts. I recommend finding a used copy. It holds up remarkably well.

The second inspiration was Meghan Duffy’s and colleagues’ recent discussion of how to organize an Intro Bio version of Ecology. I think I lifted Principle 2 and 4 directly from that blogpost. Lots of good pedagogy there.

So here is my working list of the Ten Principles of Ecology, stated first as tweet-worthy statement, followed by a short explanation of each. The idea is that my 48 students will be seeing this the first week of class and we will sample, expand on, and recombine them throughout the rest of the semester.  I realize every ecologist is different, and that this lays bare my own intellectual DNA on the subject. None-the-less, I’d love to see more lists like this.

Also, has anyone else tried a similar approach to structuring their class?  That is, start with the big picture, then backfill? I’d love to hear about it.

The Ten Principles of Ecology

1. Evolution organizes ecological systems into hierarchies.

Individual organisms combine into populations, populations combine into species, species combine into higher taxa like genera and phyla. Each can be characterized by its abundance and diversity (number of kinds) in a given ecosystem or study plot. How and why abundance and diversity vary in time and space is the basic question of ecology.

2. The sun is the ultimate source of energy for most ecosystems.

Life runs on the carbon-rich sugars produced by photosynthesis; every ecosystem’s sugar output depends on how much solar energy and precipitation it receives.

3. Organisms are chemical machines that run on energy.

The laws of chemistry and physics limit the ways each organism makes a living and provide a basic framework for ecology. The supply of chemical elements and the sugars needed to fuel their assembly into organisms limit the abundance and diversity of life.

4. Chemical nutrients cycle repeatedly while energy flows through an ecosystem.

The atoms of elements like C, N, P, and Na go back and forth from spending time in living to spending time in dead parts of an ecosystem. But the photons of solar energy can be used only once before they are lost to the universe.

5. dN/dt=B-X+I

The rate that a population’s abundance in a given area increases or decreases reflects the balance of its births, deaths, and net migration into the area. Individuals with features that improve their ability to survive (i.e., not die) and make copies of themselves will tend to increase in that population.

6. dS/dt=D-X+I

The rate that the diversity of species in an area changes reflects the balance of the number of new forms that arise, those that go extinct, and those that migrate into the area. Individuals and species that have features allowing them to survive and reproduce in a local environment will tend to persist there.

7. Organisms interact—do things to each other—in ways that influence their abundance.

Individual organisms can eat one another, compete for shared resources, and help each other survive. Each pair of species in an ecosystem can be characterized by the kind and strength of these interactions, measured as their contribution to dN/dt.

8. Ecosystems are organized into webs of interactions.

The abundance of a population is influenced by the chains of interactions that connect it to the other species in its ecosystem. This often leads to complex behavior, and a key challenge in ecology is to determine what patterns of abundance and diversity can be predicted.

9. Human populations have an outsized role in competing with, preying upon, and helping other organisms.

Humans are one of millions of species embedded in Earth’s ecosystems. The ability of humans to change the planet, abetted by our large population size and technological prowess, increases our ability to shape the biosphere’s future. Humans, through principles 1-8, are currently changing the climate, re-arranging its chemistry, decreasing populations of food, moving around its species, and decreasing its diversity.

10. Ecosystems provide essential services to human populations.

These include products like timber, fiber and food, regulating water and air quality, and cultural benefits like recreation. A key goal of ecology is to use principles 1-9 to preserve ecosystem services.


A slightly less paradoxical paradox

All organisms are built from the same recipe of 25 or so elements, and so it makes sense that as you increase the supply of those building blocks to an ecosystem, you should be able to increase the number and variety of organisms that ecosystem supports. Deserts vs. rainforests, right? In a paper just out from a terrific collaboration,  we show, it turns out, things are not quite that simple.

The background

Michael Rosenzweig

Michael Rosenzweig

In 1971, Michael Rosenzweig (MikeK’s Ph. D. advisor and friend of the Kaspari lab) used a suite of differential equation models to explore why adding nutrients to an ecosystem so often reduced, not enhanced, the number of species. He coined the term “Paradox of Enrichment” to capture this incongruity. Since then Rosenzweig and a who’s who of ecologists–particularly terrestrial botanists–have documented this Paradox and compiled a long list of mechanisms that could explain it.

A few years ago, our lab joined a team of collaborators to take on the relationship between nutrient availability and diversity in tropical soils. We did so using a grand experiment initiated by Joe Wright and colleagues at the Smithsonian Tropical Research Institute in Panama.  Since 1998, we have been fertilizing 40×40 m plots with the “big 3” nutrients–Nitrogen, Phosphorus, and Potassium–as well as a cocktail of micronutrients from Boron to Zinc. In 2012, as part of an NSF MacroSystems grant, we visited each of these plots, sampled soil and litter, and used a variety of molecular and traditional methods to count the number of species of bacteria, fungi, and invertebrates on the essentially 9 different kinds of tropical soils generated by this experiment.  I would say that our results, out early online in the journal Ecology, were surprising, but, what happens when you quantify the absence of a paradox?

Gigante Fertilization Experiment

The Gigante Fertilization Experiment, one of the largest, and longest running such planned experiments on Earth. (You can say that humans are actually doing a pretty good job of adding Nitrogen and Phosphorus and a variety of metals to Earth’s ecosystems in a less scientific, tho none-the-less deliberate manner).

The results are wonderfully complex, but here are some highlights.

Bacteria, Fungi, and Invertebrates each had their own “biogeochemical niche”.

If you plot out the magnitude of diversity responses to nutrients–positive or negative–the resulting fingerprint differed among the big three soil supertaxa. Everybody’s diversity suffered when nitrogen alone was added. This is as close as you get to a uniform Paradox of Enrichment. After that, Bacteria increased the most (and modestly) with phosphorus (P), fungi were all over the map but tend to do better with potassium (K), and invertebrates showed the biggest increases when combinations of nutrients were applied.

Why does nitrogen do a number on soil diversity?  The standard explanation is that there are a handful of weedy taxa that just thrive when nitrogen is superabundant (fertilize a prairie with urea and you wind up with tall, green patch of invasive grass). But we find little evidence for that: things that increase on N plots tend to be rare, and increase when any nutrient is added (and some of them are icky. See below).

Instead, we suggest that nitrogen, which tends to acidify the soil, sets off a chain reaction by which aluminum–a toxin to most life–leaches into the water supply and bathes the unfortunate members of the brown food web in a bath of metal. In other words, nitrogen doesn’t seem to favor a suite of nitrogen specialists, it releases an all-purpose toxin that stresses everybody out.  That’s our new working hypothesis. It needs to be tested.

Fig 1 Richness ES w treatment.JPG

A summary of our results. The Effect Size is a way of uniformly expressing the change in diversity, positive or negative ,of a fertilizer compared to unfertilized plots. For example,  fertilizers tend to have a bigger effect on invertebrates than bacteria. More and more complex fertilizer combos are arrayed toward the right. 

Combo’s of nutrients often enhance diversity more than individual nutrients.

One common pattern among plants is that if adding one nutrient, say nitrogen, drops diversity in a prairie, adding two, say nitrogen and phosphorus, does so even more. We don’t find much evidence for that. Instead, the two groups with big, complicated genomes–the fungi and invertebrates–show the largest increase in species diversity when micronutrients are added. Invertebrates do almost as well when the big three nutrients, N, P, and K, are added in tandem. What is it about nutrient combos that favor mushrooms and ants?

We suggest that one reason is that critters with large genomes need a ready supply of chemical elements to build and maintain their more complex metabolisms. It is fun to ponder such a link between an organism’s metabolic diversity–the number of enzymes it has linked together in intricate pathways (many of which require a metal like Zinc or Copper to operate properly) and its metal-craving. Again, this is a working hypothesis. But we have already shown that decomposition in this system–the combined action of the metabolisms of the brown food web–shows big increases on the +Micronutrient plots in the Gigante experiment.

Of the three soil supertaxa, the bacteria and invertebrates have the most similar biogeochemical niches.

One might have guessed that fungi and bacteria–both “microbes” that break down “detritus”*–might respond similarly to the same nutrients, but this doesn’t seem to be the case. Fertilizers with a strong effect on bacterial diversity tended to have little effect on fungi, and visa versa. On the other hand, the fertilizers that really knocked down bacterial diversity, N and NK, did the same for invertebrates.

Many soil inverts have rich microbiomes in their guts that allow them to make a living in the brown food web. Could this be one cause of their similar responses. Does the gut flora of millipedes also suffer in bacteria-poor soil?

Fig 2 Covariance of Diversity Responses

When you contrast the effect of a fertilizer on diversity between each combo of the soil supertaxa, fungi and prokaryotes (bacteria) do their own thing, while bacteria and invertebrates tend to response more similarly. 

There are weedy taxa in each of the three supertaxa

Which taxa thrive regardless of what nutrient you drop on the soil? It is somehow satisfying to find that among the invertebrates, the Blattaria (aka roaches) can’t say no to any fertilizer.

It is a little more chilling that among the fungi, only the Chytrids, a phylum that contains the infamous B. dendrobaditis, a deadly pathogen of amphibians, increases in diversity in response to almost any fertilizer.  We are fertilizing the planet. Just a pattern thus far. Someone should look into it.

Fig S1 Effect sizes by subclades

Subtaxa of the soil supertaxa that respond positively or negatively to a given fertilizer by at least one standard deviation over controls. See paper to suss out all the abrev’s.

What’s the take home?

Humans are rearranging Earth’s biogeochemistry–depleting fertile soils through erosion and dumping enormous quantities of N, P, and C and a mess of metals into the biosphere. Our study contrasted three grand hypotheses for how fertilization shapes diversity–by increasing abundance, by favoring a subset of competitive species, or by acting as toxins. Much of what we ecologists know about these nutrient effects comes from the experiments of our botanical colleagues. And those experiments consistently point to fertilizers favoring a subset of high-nutrient specialists that drive other species locally extinct. The Paradox of Enrichment is thus an inevitable result of simplifying the environment with fertilizers to favor a handful of species.

But by focusing on three soil supertaxa–the bacteria**, fungi, and invertebrates–that account for most of the diversity of life on dry land, we find evidence for some nutrients enhancing abundance, others acting as toxins and for combos of nutrients ameliorating toxic effects. For some of the reasons why, check out our Discussion. Not surprisingly, spatial scale comes into play. And how metabolisms are rejiggered.

An enduring question is how the combined diversity of life in any given ecosystem–from trees to mites to microbes–will respond to the slings and arrows of anthropocene fortune. The answer “it’s complex” shouldn’t surprise. At the same time, one path ahead, we think, lies in  contrasting how tiny organisms with effectively infinite population sizes and evolutionary potential to match respond to global change compared to their large colleagues that can be comfortably enumerated with m2 quadrats.

*Combining fungi with bacteria** as “microbial” is akin to combining petri dishes and planets as “matter-intensive phenomena”.  Likewise “detritus” can be anything from a dead leaf to a dead armadillo. *Celebrate* the diversity.

**Again, I beg forgiveness from microbial ecologists for using the shorthand “bacteria” or “prokaryotes” to lump together the archaea and eubacteria. I was raised by ornithologists.

A short essay on Koechner’s Criterion and its relationship to success in Grad School

During a layover at DFW airport, I opened Twitter to find a lively discussion spurred on by Terry McGlynn, @hormiga, who (and I paraphrase) encouraged grad students to seek mentors who encouraged living normal lives, using the specific example that Saturday work days should not be mandatory.

The thread quickly expanded into one about work/life balance in academia, many of the arguments and counterarguments (in the form of tweets and counter-tweets) appeared to hang on definitions both of success and what “normal” looks like or should look like in a better world. All of it had the subtext that bubbles under many such discussions—“Am I working hard enough?”. In my experience, there are few more troubling questions to an academic scientist, or any creative person.

I dedicated a blog to this and related subjects in the (my gawd) late 2000’s called “Getting Things Done in Academia” (rebranded “Survive and Thrive in Grad School”) that is moribund but still out there, and hopefully moderately useful.

So there I was, sitting in the airport, waiting for my flight to OKC, and reading tweets and counter-tweets on an important subject that I obsess about. What was I to do?  I tweeted the longest thread of my TwitterCareer, I think. Three whole tweets. I started with the proviso that Twitter is a mixed bag for holding such discussions (Boy Howdy, nothing like leading with understatement). Then, because I found it an intense distillation of one view, I quoted EO Wilson’s comment from Advice to a Young Scientist, that said for a real scientist, every vacation should be a “working vacation”. I ended with my thoughts on that key question “How hard should I work as a grad student/academic?”.

I thought I’d add a few observations. In the spirit of Twitter, they will not be organized into a coherent whole ;-). I suppose I could blast out a thread, but it is easier to get some thoughts down in one place, as a blogpost, and link to them. (I wonder if such a stratagem, once things get hot, might be a productive way to carry on the discussion. As good as Twitter is for starting a discussion, I would rather read one continuous argument).

Assumptions and Definitions

1) The best definition of a successful life, that I’ve yet heard at least, is from the comedian David Koechner, @DavidKoechner, who said

“If you are doing what you love and have someone to share it with, congratulations, you are a winner.

I think this should be called Koechner’s Criterion. I mean, does it set up a discussion of work-life balance, or what?

2) Scientists are scientists because they are intensely curious about how Nature works, and want to discover new stuff and share those discoveries with others, particularly those that are curious about the same stuff. The most successful scientists, however we define them, do this discovery/sharing stuff more and better than others. They also have a higher probability of getting jobs where they are paid to do science.

3) We live in a stochastic universe where sheer dumb random luck can be important and bad luck is rampant.

4) There are skills,  mental attitudes, and societal tweeks that, once achieved, improve our chance of being successful.

5) We spend much of our lives pursuing 1, desiring 2 as part of 1, fearing 3, and seeking 4.


1) There are as many formulae to being a scientist (or artist, teacher, craftsperson) as there are people. But I think that all routes to mastery include a passion for the subject, an aptitude for it, and a desire to create the opportunities that will allow one to develop and prosper. For many people, this leads to a more or less single-minded devotion early in their careers to master a difficult task. However, the diversity of humankind guarantees that there are other folks whose sheer brilliance will allow them to succeed without breaking a sweat. There are still others who are amazingly effective, doing a lot with less time, because of, not in spite of, their strong relationships. Against that diversity of approaches all are judged by a common outcome: the discovery of new things and sharing them with an interested audience.

2) There are also a lot of ways folks decide to share their lives together (including living alone), and the best institutions honor that variety. Family leave, maternity leave, equal pay for equal work, are all hallmarks of a civilized society, in no small part because they help each person fulfill their potential.

3) The happiest biology departments I know find a way of of making steady progress toward maximizing Koechner’s Criterion for their faculty, post docs, staff, and students.

Conclusions in the form of suggestions to Grad Students

1) A key task in a young scientist’s career is to find a mentor who is the best match for the way they seek to develop their career and maximize Koechner’s Criterion. The single best way to find that match is to talk to the graduate students and post docs of possible mentors.

2) It is an old, but true adage that its not just the time you put it, but how you use that time (i.e., working hard versus working smart). If you haven’t already done so, read these two books: The Seven Habits of Highly Effective People by Stephen Covey, and Getting Things Done by David Allen. Seven Habits focuses on the strategy of designing a creative life, GTD is about tactics. There’s work-life balance baked into both of them.

3) Unplug. Daydream. Daily. One key element of Zen Meditation is to calm the Monkey Mind. Twitter and Facebook are the Monkey Mind incarnate. Your best ideas will come to you when you release yourself from the data stream. That also means your best ideas will likely come to you on vacation, so don’t forget to bring a notebook.

4) One good way to center yourself and set your own expectations is to read memoirs of scientists. Sapolsky’s A Primate’s Memoir, Fortey’s Trilobite, and, of course, Jahren’s Lab Girl are all awesome and different.

5) I would be remiss if I didn’t conclude this essay by plugging EO Wilson’s Advice to a Young Scientist. Since when does one read an advice book in which you agree with everything in it? The book presents a series of chapters that have got a lot of discussion going  (one review). Find a new or used copy, or check it out from the library. Or start with Wilson’s Ted Talk on the subject.

Herbivores like a little salt with their protein

One of the questions that drives a lot of our work is “Of the 25 elements required to build organisms, how many of them help regulate the abundance, activity, and diversity of organisms in ecosystems?”. In a freely available paper just out as a Report in Ecology, we provide a framework for the many ways that nutrients can interact, then test how Sodium (an AntLab favorite) and Nitrogen+Phosphorus (the preferred pair of the “Stoichiometric-Set”) interact to shape the abundance of the invertebrates above- and below-ground.

We were particularly keen on seeing if Na acts as a catalyst, increasing an insect’s ability to efficiently convert Nitrogen and Phosphorus into more insects, or if the two sets of nutrients acted more or less independently.


Four possible ways that two sets of nutrients can interact. We were betting on Serial C0-limitation, where Sodium catalyzes access to N and P. Didn’t always turn out that way.

The experiment was pretty classic field ecology: we furnished m2 plots with either water, N+P in the quantities used by the NutNet experiment, 1% NaCl solution, or both. We measured the soil and plant responses, and bugvac’ed the above-ground invertebrates with a modified leafblower (see picture above). For below-ground invertebrates, we used a very messy, but unfortunately best method available, flotation extraction of soil cores from the center of each plot.


A soil core from one of the experimental plots. Somewhere in there lurk oribatids, collembola, and other tiny inverts.

As an aside, as someone who cut his ecological teeth in the Nebraska Sandhills for his undergrad and master’s work at the University of Nebraska (see below), and has spent a good fraction of my days since then in the tropics, I am very much enjoying spending time in grasslands again. An NSF project with Nate Sanders on the role biogeochemistry plays in grassland food webs will keep me in the grass for the next few years.


A 21-year version of Mike Kaspari above WhiteTail Creek in the Sandhills of Nebraska. Note the Nikon K-100 camera, and the spiffy digital watch. And yes, those are cutoff jeans. It was a different time then.

After the sampling came the microscope work. Leafhoppers are by far the most common insects in the aboveground samples, and their mobility and subsequent ability to respond quickly to manipulations is something we want to investigate further. As predators go, spiders were common and diverse, and clerids were the most common beetle group.


Looking forward to getting to know the leafhoppers, a group whose English name is simple, and whose latinate name defies easy memorization.



One of the big questions going forward is how quickly predators accumulate on plots that first attracted their prey. The spiders were dauntingly diverse on this patch of Oklahoma prairie.



One of the greatest pleasures of invertebrate microscope work is getting acquainted with taxa like these clerid beetles that are common, but that I’ve never had the occasion to look at before.

Results: Both sodium and N+P shape both communities, but in different ways

In the grass and forbs above-ground, all three treatment plots differed from controls. Adding salt increased the abundance of insects, and adding N+P increased it even more so. However, these effects seemed to be independent of each other (strike one for the Na as a catalyst hypothesis). Since plant height and mass increased on N+P plots and not on Na plots (suggesting that while NP was a nutrient for the plants, adding Na had no negative effect on the first trophic level), we think that NP provides a double bonus of more food, and more habitat, while Na just provides the food. We will test this idea more this coming summer.

Below-ground, however, was a different story. Here Na seemed to act like a catalyst. Alone it had no effect on the oribatids and collembola that dominated belowground. But combined with NP, it boosted the modest increase of abundance where NP was added by itself. Here, our working hypothesis is that NP fertilization increases the food supply, and thus increases the demand for Na. We are keen to follow this up this summer as well.

So there it is, so far. The plans are to expand both the kinds of experiments and their distribution, to add experiments farther north in more Na-deprived grasslands. One project has a particular appeal: as NP + NaCl is roughly the recipe for urine, and urine is the single most effective way to fertilize a patch of prairie, how do community dynamics respond to a splash of Bison Piss?  Is there a predictable succession?  And how do the hundreds of species find, exploit, and deplete this resource? What are the patterns of succession?  Lots to learn.


When the science news is bad, write your local paper


Recent events have found me struggling to find some way to be useful. I decided that one thing I could do was to write–on a regular basis– a letter to the editor of the The Norman Transcript, our local paper. The bad news from Crowther et al in Nature served as the grist for this week’s letter. Writing it scratched a number of itches: it gave me a shot at explaining negative and positive feedback, it provided readers with actual phone numbers (not just the link in this version) to phone the local offices of our Senator’s and Representative, and it pressed the notion that there is always hope when people engage, even when times are dark. Below is a slightly modified version of the letter.

To the Editor:

Last week was a bad one for Planet Earth. While her workings are pretty complex, it has long been evident that when we pump millennia-old carbon into the atmosphere we warm the planet. Many of us scientists had hoped that this warming would be slowed by a variety of feedbacks. Trees, after all, scrub CO2 from the atmosphere; more gets sucked up by our oceans. That, we hoped, may buy us more time to move away from fossil fuels and build the solar panels and windmills that are springing up over the state. These natural feedbacks protect us in the same way that, when you notice a bridge is out, you step on the brakes long before catastrophe strikes.

Well, there is another kind of feedback, and for that we can thank the microbes. Boatloads of carbon rest in the cold earth. And, just as we keep our bacon in the fridge to keep it from rotting, that cold earth has kept its microbes from chewing through the countless tons of dead carbon just below our feet. Now, 49 scientists have together revealed in the journal Nature that the warming Earth is waking the microbes from their cold torpor. And they are hungry. As the microbes break down the soil CO2 pours into our atmosphere. This warms the Earth and makes the microbes even hungrier…

You are, by now, beginning to see how this other feedback works. It is as if, back in the car, you stomp on the brake pedal. Or…what you *thought* was the brake pedal. The car lurches forward, racing toward the chasm. So you stomp harder! (Why aren’t these brakes working??) In the same way that the car accelerates toward the chasm, every fraction of a degree that we further warm the planet drives Earth’s microbes to empty one of its last storehouses of carbon, making the problem even worse.

Bad timing, right? Just as we should be redoubling our efforts to find a solution, prez-elect Trump calls climate change “bunk” and his advisor on NASA wants to delete such “politicized science” from its budget. This is code for the hundreds of satellites that watch our Earth, informing our farmers about soil moisture, our sailors about sea ice, our friends and neighbors about approaching storms. All because some of that data is used by scientists to diagnose our warming planet. It is bad enough our car is hurtling toward a cliff. Trump’s NASA policy would disable the speed gauge and paint over the windows.

Now more than ever we need to phone our Senators and Representative. We need to urge them to preserve NASA’s world class Earth Observatory program. Why do I think you and I can convince climate change deniers that we need to do more, not less? Because over the past 23 years at OU I have had the privilege of teaching science to thousands of Oklahomans. These young citizens see the evidence, and in our conversations (and in anonymized polling) overwhelmingly agree humans are warming the Earth and that there is still time to save it. Full stop. These same people are our future governors and legislators, will be paying taxes when some of us are retirees, and are thinking about raising families of their own. I suspect these are some of the same people who will be answering the phone when you make those calls. They are our future. They are our hope.

Mike Kaspari
Norman Oklahoma

One way to build a dietary generalist

By Karl Roeder

We are incredibly excited to announce that our work on fire ants and isotopes has just been accepted in Ecology! The work primarily revolves around understanding trophic variation across a population of one of the model organisms of myrmecology: the red imported fire ant, Solenopsis invicta. Essentially, we are looking at where you rank in a food web and what did you eat to get there. To expand upon this, we had been thinking about ways to understand the how, what, why, and where of stable isotope variation that regularly occurs in the published literature. Stable isotope analyses have increased in their use in ecological studies as a tool where nitrogen (d15N) can be used to understand trophic position/structure and carbon (d13C) can be used to map out where the dietary source came from, in this case C3 or C4 plants.

Despite a lot of myrmecologists loving morphological measurements, it seems few had tested how size, both body and colony, may affect trophic variation within a species. A question that we were intrigued with. Perhaps even more surprising was a real lack of good information at the colony level, as most stable isotope studies with ants focus on differences across species in an assemblage or community. We set out to answer these question by measuring a variety of different sized workers and colonies across multiple time periods throughout a year in a small 0.5-hectare old field in Oklahoma. To our surprise, we found an unprecedented range of values occurring within a population. A range that was comparable to whole ant assemblages from prairies to tropical rainforests. The red imported fire ant, while acting as a generalist at the species level, may in fact be specializing on particular dietary items at the colony level.

While there are still mechanisms to work out, we believe this is a wonderful step towards disentangling the trophic ecology of the red imported fire ant and potentially other invasive species. Given the wide diet breadth we observed in such a small area, we hope this will encourage researches to think more about the natural history of their study species, and combine this information with physiological measurements to really attack questions surrounding their biology.

This work was generously funded both by the National Science Foundation and a University of Oklahoma Biological Station summer fellowship.


What journal gets the first peek at your manuscript? Results from a year of ruminating.

For the past year or so I’ve been ruminating about a problem common to academics: Once you have completed a MS, where do you send it? And how do you follow that process through to completion (i.e., publication)? The literature explosion, as well as the increase in venues and ways to publish, has made this topic an acute one.

What follows is my provisional conclusions from these ruminations: Best Practices V1.0. These practices arise, hopefully somewhat logically, from some axioms. Both have been particularly shaped by discussions with Nate Sanders, by a terrific essay by Brian McGill and by multiple tweets, blogposts, and discussions with Ethan White.

That said, I strongly suspect they will disagree with some of what follows. But then, the more I work on this problem, the more I see that its solution lies somewhere within the remarkable g’mish of practicality, laziness, efficacy, ambition, fear of mortality, and ethics that characterizes doing science nowadays. I, like you gentle reader, am just trying to figure out what works. And caveat emptor: all that follows arises from the consideration that I am a mid-career professor. I suspect much of this will ring true to most. But I hope to see how other folks–grad students, post docs, junior professors, folks at undergrad institutions–formulate their own best practices.

My Motivator

Ecologist’s die with 99% of what they have learned between their ears.” Dan Janzen.

I was a sophomore at a faculty reception for Janzen at the University of Nebraska when he made this offhand remark. It has never been that far from my thoughts since. It came to a head in the past couple of years,  when I found myself spending too much time revising papers rather than working up data. As a consequence I was falling increasingly behind. This is bad news for any number of reasons: 1) somebody paid for those studies;  2) when they paid for it they implicitly contracted me to analyze it (not just dump it into the grey literature); and, 3) I really want/need to know the answers–out of sheer curiosity and my desire to move our science a wee bit forward. Which led me to spend some time, in quiet moments, over emails, and over beers, coming up with

My Axioms

1. The biggest improvement in a MS happens from Draft 1 to Draft 2.

That improvement is maximized when Draft 1 marinates for a month or more. The process of cranking out Draft 1 wears a lot of ruts in the way I think about my question, the presentation, the results, and their interpretation. A one month hiatus allows my subconscious to work on the problem; at the odd hour, I find myself scribbling notes on various and sundry ways to fix things in the MS. That hiatus (sometimes, but not always a deliberate one in the past) has always allowed me to return with a fresh view and significant improvements.

2. Outside reviewers improve manuscripts in a qualitatively different way than the hiatus between Draft 1 and 2.

Peer reviews are gold. Peer reviewers have the useful feature of existing outside my head. They recognize flaws to which I am blind, point out weak logic that I try to force through by dint of will, and highlight when I communicate poorly (default mode: if a peer reviewer misunderstood something, I didn’t communicate it well). The availability of good peer reviews–like sufficient time, good collaborators, field station happy hours, and grant money—improves the quality of our work. Access to peer reviews should be celebrated but conserved.

Yes, of course some of them are jerks.

3. The rate of manuscript improvement is a positive decelerating function of the number of reviews.

As you work to incorporate the advice of reviewers, a fairly predictable thing happens. Subsequent rounds of reviews become less useful (even if the average length of those reviews is remarkably conserved). In short, there is a rapid saturation in quality of your manuscript with repeated exposure to peer review. In the worst-case scenario, tangible improvement gives way to the stochastic chasing of reviewer whim.


Time spent revising a manuscript means time taken away from other creative endeavors.

4. Papers are not read if people don’t know about them.


If a potential reader does see your intriguing title and abstract on a journal website, they should not have roadblocks placed between them and the rest of the paper.

5. Scientific societies are natural places for scientists to aggregate, discuss, train, synthesize, and publicize the work they celebrate.

The flagship journal of your scientific society is a go-to place for those interested in your kind of work.


Money invested in your society, including publication fees, is likely to go to the people and activities you believe in.

OK, we have our axioms. What do we conclude?  Here is my provisional rules of the publishing road for the next few years.

Kaspari’s Best Practices for Submitting Papers V1.0

1. After completing a MS, let it sit for at least one month.

Then open it with fresh eyes and revise as needed. Then send it for friendly review. Then, and only then, send it out to a journal, and exploit the peer review system.

2. Send the manuscript to a society journal that best matches its audience

Here is my current list:

PNAS for the very best stuff of widest interest (Bert Hoelldobler told me “PNAS is where you go when you’ve got a great story and want the space to tell it properly.”.

Next is Ecology and American Naturalist.

Next is Ecography, Functional Ecology, BiotropicaEcological Entomology, and Soil Biology and Biochemistry.

3. If a paper is rejected, do not “shop it around”.

Get it out. Ecosphere and Ecology and Evolution are two open access online journals from societies I believe in.

4. Maximize access

Not all society journals are open access. We are not where we want to be, but the times they are a changing. Be an agent of change. That said…

5. Publicize your work

Get the word out. Email PDFs to colleagues who should be interested. Given the publication glut–and my general sense of the universe’s spiraling, magnifying disarray–I am delighted when someone thinks of me enough to send a PDF they think I should know about. And it happens remarkably rarely.

Moreover, if you’ve spent all that time discovering something new and interesting, then

Tweet it

Blog about it

Post links to the paper on your website (when you can inkway, inkway)

Send your University Public Relations your pithy public summaries from social media.

Every tweet and blogpost is vital practice at communicating our science to a broad audience.


How will I know in five years if Best Practices V1.0 has been a success?

  • No manuscript will remain a manuscript more than 1 year (a paper is no longer a manuscript once it is accepted/in print).
  • I will spend no more than 30% of my creative energy revising manuscripts.

Until then.