The Kaspari Lab

Not all male ants are “sperm missiles”

Image from the phenomenal  Alex Wild.

In a new review Juergen Heinze discusses the adaptive nature of male lifespans in ants. Far from being mere short-lived “sperm missiles”, in EO Wilson’s words, many ants, particularly in the tropics, live an independent existence, sipping nectar and looking for mating opportunities.

Juergen highlights the contributions of  AntLab alum Jon Shik, particularly his  Life History Continuum Hypothesis.  When most folks think of mating ants–and you are likely thinking about them right now–they picture swarms of males and females in some weird, lascivious mating scrum. Kind of a cross between “Dirty Dancing” and “Mad Max Beyond the Thunderdome”. Such scrums leave the wreckage of spent males littering the landscape, typically to be picked up by other ants and carried off for food. Ahh, the cycle of life.

Jon builds the case that “female calling”, the ancestral mating system in ants, favors males that can endure life longer outside the nest. In female calling, new queens emerge from their natal colony and release a pheromone to lure in male suiters. If no male comes along, they may return to the nest and try again the next night. In this slowed down breeding system,Jon argues, it is in the male’s best interest to be persistent in the search for his elusive one-on-one rendezvous with a female. Species with colonies that subscribe to female calling, he shows, provision their males for the potentially long series of nights ahead.

Considering how important the relatively short-lived sexual window is in the long-term existence of a typical colony, we know surprisingly little about male biology, or even the phenology–the timing–of ant flights. Early in my career, I gained access to hundreds of vials from Light Traps and Malaise Traps on Barro Colorado  Island, and spent many delightful hours sorting the winged queens and male to species. My ultimate goal was to test the hypothesis that the some 400 species of ants on the island may partition the calendar, each flying in their own designated window.

Screen Shot 2016-06-10 at 11.02.55 AM

The number of ant queens and males, two species from four genera, flying across the lunar months on Barro Colorado Island, Panama. Even Gnamptogenys hartmani, which prefers to fly in May, will dribble out alates over most of the year. Many of these species exhibit the mating syndrome “Female Calling”, where queens lure individual males using pheromones.

I was shocked, shocked (!) to find my hypothesis pretty much destroyed by something even cooler–that a large fraction of tropical ant species fly year-round. Jon’s work nailed the connection to their mating system.

This still leaves a lot of potential for good work. To me, one of the big questions, relevant to any group with lotsa species in one place, is how do they divy up all that reproductive bandwidth?  Assume, for example, that queens of 200 or so species in the forest of BCI may be releasing their pheromones on a given night, and all those different molecules swirl through the still humid night air. That’s a lot of perfume to sort out. No wonder males are long-lived. They have to be patient.



On the salutary effect of the Kudos Email

If you like someone’s work, don’t just cite them, write them!

We scientists are hard on ourselves. We obsessively check and recheck our data. We expose our nascent manuscripts to the keen scrutiny of co-authors, mentors, and anonymous peer review. Most of the time, the reviews are critical. Sometimes they are downright unkind. Ultimately, we hope, our revised manuscript pasts muster. Then we get to enjoy the scrutiny of a copy-editor and in the process discover typos that have survived, unscathed, through the gauntlet. Little wonder that on publication our feeling is mostly relief, rarely joy.

Why do we do it? As scientists, we constantly court criticism because it helps us to find our errors. I rarely read a review that is not in some way constructive, if only in the sense of “I need to communicate this better”. (1) And in any creative endeavor, discovering our failures, large and small, improves our work in the short run and builds our scientific intuition, project by project, review by review.

With this constant barrage of criticism, it’s not surprising that Imposter Syndrome (2)–the inability to internalize success while living in a constant fear of being exposed as a fraud–is so rampant in science. It is common in every creative endeavor. And don’t make the mistake of thinking that Imposter Syndrome is limited to novices. Even the most alpha scientist at a conference is only two perceived slights away from re-living their first year of grad school.

A modest suggestion: the Kudos Email

We occasionally come across a paper that inspires us. It could be a citation classic, an obscure paper that has flown under the radar, or something new that you discover as part of your regular reading routine.

That paper’s author deserves a Kudos Email. (3)

A Kudos Email communicates to the author that you enjoyed and were inspired by their work. It includes something concrete to back up the praise: “Someone finally figured out how to experimentally test X in system Y”, or “Figure 2 has changed the way I think about X”, or “I’ve passed this onto my reading group”, or “I’m looking forward to using your paper in my class.” (4). A Kudos Email arriving on the desk of the harried scientist (especially if is from someone they don’t know) can do a lot to balance the psychological trauma of Reviewer 2. It is also a pleasant and effective way to introduce yourself to someone you might want to get to know.

I have been writing Kudos Emails for years. They are easy to compose because they form in your brain as you read the paper: “Wow, that is really cool.” ,”Yep, gotta try that out in my system.”, “I can’t wait to tell my lab about this one”. Kudos Emails don’t have to be more than one short paragraph. And here’s the great part: everybody–from grad students to proto-emeritae–likes to get Kudos Emails. Everybody likes to hear from someone they don’t know telling them they did good. In my experience, the most common response to a Kudos Email from a young scientist is “Thank you very much! I’m very happy with the work and am glad you liked it.”. A common response from older colleagues:  “Odd. Nobody’s every written me an email like this before.”. (5)

Here’s the thing. There is more than one way to increase the quality of work in our beloved science. Sure, a key way will continue to consist of writing reviews that, even done concisely and with compassion, still sting a little. Why not also praise published work we admire, and do it while the paper is fresh in our minds? Why wait for the author to tally her citations? Investing in some Kudos Emails encourages the very scientists that you think are doing good work.

There is also a fringe benefit to Kudos Emails. When we express gratitude to our colleagues for teaching us something new, it can do wonders for our own outlook (6).

We’re all in this together.

(1) And even reviews by cranky old assholes teach one the very important lesson “Don’t be a cranky old asshole.”.

(2) Just Google it.

(3) From the online Oxford Dictionary. Usage: Kudos comes from Greek and means ‘praise’. Despite appearances, it is not a plural form. This means that there is no singular form kudo and that the use of kudos as a plural, as in the following sentence, is incorrecthe received many kudos for his work (correct use is he received much kudos for his work).

(4) The “one concrete thing” rule ensures the by-now-flabbergasted recipient that you are sincere and not just trying to get them to submit their next paper to your new open access journal “Paradigms in Science Capitalization”.

(5) Something like this used to happen–kind of–in the bad old days of paper reprints. Folks like me would carry pre-printed postcards to the library, skim the new journals, and when we saw something we liked, scribble out the particulars on the back of the card, mail it to the author with perhaps scribbling a hand-drawn heart and “Thanks!”, and then wait for a copy of the reprint to arrive in the post. Getting reprint requests communicated that *someone* was interested in your latest work.

(6) Choose to be grateful by Arthur Brooks, New York Times, 2015

On writing a strong NSF pre-proposal

Since 2013, NSF DEB has used a two-step merit review system–a 5 page pre-proposal, followed by a 15 page full proposal–to inform the process of awarding grants. The full proposal had long been standard. The pre-proposal, just four years old, is still relatively new. I have served on three DEB pre-proposal panels. Over the course of that service I have made some observations and formed some opinions as to what makes for a successful pre-proposal. I’d like to share some of those observations here.

As panel work is anonymous, I scrub any reference to specific panels, panel members, or program officers. My intent is to walk you through the process of being picked for a panel, writing reviews, and the panel work itself, adding some observations and suggestions where appropriate. I then step back with some conclusions about why some pre-proposals are invited for a 15-page full proposal, and others aren’t. I finish with some suggestions as to how best to craft a strong pre-proposal.

Caveat emptor: these are my opinions. To paraphrase Anne Elk these impressions that I have, that is to say, which are mine, are mine.

First some data

DEBrief ‘s recent post included a graph that summarized success rates versus average scores for both pre-proposals and full proposals. The x-axis shows the average score of the reviews (n=3 for pre-proposals, n>3 for full proposals), where a reviewer score of Poor=1, Fair=2, Good=3, Very Good=4, and Excellent=5 (more on that scoring system below). One clear lesson is that the key challenge for pre-proposals is to achieve an average score of ca. 4 or above. That is, scores dominated by VG’s and E’s.


To see how, let’s first examine how the proposals are reviewed.

Getting selected for a pre-proposal panel

Around the beginning of the year NSF emails a bunch of folks asking if they would be available to serve on a pre-proposal panel, noting that at this point they are just establishing a pool of potential panelists. You are asked if you are interested. After replying in the affirmative, a subsequent email asks which of three panel dates fit your schedule. The next email about a week later says “Thank you serving on a panel service; please fill out these forms, and arrange hotel and airfare through our designated travel agency.”

Observation: Note NSF pays a stipend for this work, out of which you pay for your hotel in Arlington (not cheap) plus airfare, and expenses. Note also that the following January you will receive an IRS W9 for this stipend, which means it’s miscellaneous income, which means that if you are reasonably prudent, after paying some of this stipend back to Uncle Sam as income tax, you will about break even financially when serving on an NSF panel.

Identifying the pre-proposals you would like to see

Soon thereafter you are emailed a list of ca. 150 pre-proposals listed by title, Principle Investigator, and the PI’s affiliation. You are asked to speedily 1) identify any conflict of interests that weren’t already apparent from your working address; and, 2) assign each pre-proposal into one of three categories: effectively “1. I would like to review”, “2. I’m OK with reviewing”, and “3. I don’t want to review”. This process is a little mind-bending, and takes about and hour and a half.

Observation: The title of your pre-proposal is important. If you want to give the Program Officers at NSF their first, positive clue as to how the scientific community views your project, craft a title that interests non-specialists in your field. The flipside is just as true: if you want to send the opposite signal, load your title up with jargon and make it obtuse enough that you earn loads of “3”s. The POs will then have to work to scrape together three panelists. Some practical advice (which also applies to manuscripts): craft 5 titles for your pre-proposal and market test them–in your lab or among your colleagues–to find which one sparks the most interest. If you are like me, you will be surprised as to how often the consensus is not your favorite.

Reviewing the Pre-Proposals

About one month before your panel meets, you are sent a list of 20 or so pre-proposals to review. They consist of a summary page, a personnel page, and a 4-page project description.


NSF instructs reviewers to consider the following criteria sequentially when judging the quality of a pre-proposal.

A. Is there a clear and compelling question(s) of general interest to the field?

B. Is the question(s) well motivated and justified within a broader conceptual framework?

C. Does the experimental design/approach logically and feasibly address the question(s) posed?

D. Are the senior personnel qualified to conduct this research?

E. Is there a credible plan for broader impacts?


NSF’s instructs reviewers to integrate these criteria and generate a score from the following menu. It is OK to include two scores (i.e., VG/E, F/G). The scores and their official NSF meaning:

Excellent–Outstanding proposal in all respects; deserves highest priority for support.

Very Good–High quality proposal in nearly all respects; should be supported if at all possible.

Good— A quality proposal, worthy of support.

Fair— Proposal lacking in one or more critical aspects; key issues need to be addressed.

Poor— Proposal has serious deficiencies.

The challenge for reviewers

Three reviewers, all members of the panel that meets in Arlington, are assigned to each pre-proposal. There are no outside, ad hoc reviewers on pre-proposal panels. It is my experience that the NSF program officers pretty much unfailingly assemble a panel who take this job very seriously, and who, collectively, represent the field the NSF subprogram is tasked to serve.

Each panelist has one month to review and write substantive comments on ca. 20 proposals; on about a third of those, you are expected to lead the discussion and write up the final panel summary. This service is squeezed into the usual academic spring schedule: the student committee meetings, job interviews, field season planning, and general end of academic year chaos. Luckily (or unluckily, as the case may be) you can always dip into Spring Break to finish your reviewing obligations.

As a panelist, you know that at least one Program Officer will read you review, as will the two panelists who are also assigned that pre-proposal. Unlike ad hoc, anonymous reviews, I have rarely encountered a panelist review that was not thorough and helpful. (Think of the difference between anonymous comments on social media versus face-to-face discussions). 

Observations: Most reviewers read the Project Summary and Project Description together in one sitting and then power through their review. In my experience, that typically means 2 hours of concentrated work. In that time you must grok the project, often on a topic well outside you area of expertise, judge its importance, and write a constructive review. The chief task of a pre-proposal review is to detail the strengths and weaknesses of the proposed work so as to improve subsequent efforts by the PI, whether they involve an invited full proposal or not.

Just to emphasize, you as a PI will be lucky to have one person from your specialty among the three scientists tasked with reviewing your proposal. I will return to this later, but suffice to say that the first task of a PI is to craft pre-proposal’s entry point into the problem. This entry point must entice three people and a program officer from an audience as broad as the field served by your panel. If you have some chops as a teacher, this will serve you well in writing a pre-proposal.

The Project Summary of your pre-proposal is key. It has to sing. By its conclusion, the reviewer, with a folder full of pre-proposals still to review, will already be taking notes and beginning the intellectual process of figuring out what score to assign. After reading the Project Summary, you want that reviewer on your side. This leads to another small but useful bit of advice.

DON’T copy and paste paragraphs from your Project Description into your Project Summary (or visa versa). At best, it looks a little lazy, and at worst it denies you precious space to explain in greater detail or in a complementary way what you propose to do and why. Again, the reviewer is reading this in one sitting, and, unlike a 15-page full proposal, can comfortably keep the whole thing in her head. The Project Summary and Project Description should complement and reinforce each other, not parrot each other.

Panel service

One month later you fly to DC on a Tuesday, take the metro to Arlington, check into your hotel, and go out for a beer and dinner with a colleague or two. There are a number of benefits to panel work. The biggies: you are helping in a critical communal enterprise, and you learn an enormous amount about the nuts and bolts of NSF. There are also two more fringe benefits. First, for a foodie, Arlington ain’t half bad, especially now that food trucks park near the NSF building (though I suggest you avoid the fusion truck that serves the Borscht BBQ Burrito). Second, you see old friends, make new friends, and invariably are introduced to folks–serving on the same panel–whose work you’ve long admired.

The panel process goes something like this. A long rectangle constructed from 4 tables dominates a conference room. One side of the room has a steady supply of not entirely unhealthy food and coffee; opposite is a plate glass window. Twenty-five or so panelists sit on three sides of the rectangle; at the front of the room sit five program officers. Three administrative assistants work the back of the room. Our collective job for the next 2.5 days is to caucus, decide, present, and write up summaries of ca. 150 proposals, with a broad guideline of recommending 25% for Invites for full proposal. As panelists, our primary job is to write constructive reviews that will help anybody who submitted a proposal to get a sense of the strengths, weaknesses, and how to improve their work.

The three panelists assigned each proposal, have already uploaded their reviews to Fastlane. One panelist, the “scribe”, typically gathers the other two outside the conference room to discuss their scores and decide on a ranking of “Do Not Invite” or “Invite” for full proposal. Those discussions can be relatively brief if everyone ranked a proposal “F”. They can go on for half an hour if there is strong disagreement or if everybody liked the proposal. In the former case, the trio needs to come to a decision by talking it out (in all but handful of proposals, the three reach a consensus). In the latter case, all three are interested in seeing the proposal succeed and spend that extra time making as strong a case as possible for presentation to the full panel.

Back at the table, the Program Officers hold court, bringing up proposal after proposal for discussion (after first temporarily exiling panelists with conflicts of interests). Multiple program officers are typically taking notes during the discussions. For each proposal, the scribe presents a short summary of the group’s conclusions, with interested parties around the table listening in. The other two members of the trio add their two cents. Sometimes another panel member may do so as well. The Program Officers ask a question or two, and the “Invite” or “Do Not Invite” recommendation is added to the big screen.

That done, it is up to the scribe to summarize the panel’s recommendation, using NSF’s Fastlane Panel System. The goal is to constructively identify the strengths and the weaknesses of the proposal, highlighting the consensus points of the three reviewers, as well as any new insights that arose as a result of panel deliberations. NSF is freakishly obsessed with quality control at this point–Fastlane allows the two other reviewers to comment, and the panel summary is subsequently tweaked; an amazing set of administrative assistants wordsmith further; the summary is further tweaked; a Program Officer then edits and makes queries and edits. By the time the scribe “Formally Submits” the panel summary to Fastlane, it has seen at least four editors.

Then it is up to the Program Officers who use the reviews, panel summary,  their notes, the history of the proposal, and other criteria (Hello, first time investigators!) to decide which PIs get invited to submit full proposals. The panel, whose work is now done, doesn’t see this part.

Observation: One of the strengths of NSF’s approach is that everybody who formats their proposal correctly and submits it on time earns the time and efforts of the panel. In that sense, NSF’s process is scrupulously egalitarian. However, one consequence is that, while the panel may end up classifying 25% of the pre-proposals as “Invite”, many, many of those in the “Do Not Invite” category sample the “P, PF, F, FG, G” end of the gradient (see the DEBrief post above). Keep that in mind when funding percentages are discussed. Sure, U.S. science is underfunded. Those numbers are disturbing enough. But although only 1 in 4 or so pre-proposals may advance to “Invite”, at least another 1 in 4 were not that competitive in the first place.

How pre-proposals make the cut

Let’s review the criteria set out by NSF for evaluating pre-proposals:

A. Is there a clear and compelling question(s) of general interest to the field?

B. Is the question(s) well motivated and justified within a broader conceptual framework?

C. Does the experimental design/approach logically and feasibly address the question(s) posed?

D. Are the senior personnel qualified to conduct this research?

E. Is there a credible plan for broader impacts?

Based on my observations of fellow panelists, this is how I translate the scoring system, incorporating the above criteria.

Excellent (Rare): Wow, I really want to hear what the PIs find out, and they clearly have the chops to do it. This is as close to a transformational proposal as I’ve seen.

Very Good (Uncommon): Solid all around, OR an Excellent for which B and C need strengthening.

Good (Common): Nothing really wrong with it but suffers in comparison with those receiving an E or VG. Often a weak A and B, or a great A or B.

Fair (Uncommon): A and/or B missing, OR A or B present but C is an absolute disaster.

Poor (Rare): Aggressively bad: A and B missing.

Summing it all up:

The Probability of an “Invite” is proportional to
(A2 + B2)*(C+D+E).

That is, a strong A and B followed by a competent C, D and E are the ingredients of successful pre-proposal. Given that, let’s deconstruct criteria A through E.

A. Big question? Is there a clear and compelling question(s) of general interest to the field?

There are three key words here.

Clear–The question, pitched thoughtfully, needs to show up in the first three lines of the Project Summary, and the first half page of the Project Description. You are entering P/F/G territory if the three panelists can’t agree on your question.

Compelling–This is where the magic lies, the difference between an ecology course taught by a competent and a master teacher. When your panelists caucus, they agree, “we need to know the answer to this”.

General Interest–The three panelists assigned your proposal will include, if you are lucky, one person who would be giving a talk in the same Oral Presentation as you at an international meeting. That is to say, the panelists represent a diversity of expertise, and likely two of them don’t eat/breathe/sleep your interest. Your project must be pitched to appeal to, and be understood by, any good Ecologist/Evolutionary Biologist. Imagine you study the community ecology of microbial biofilms: how would you explain what motivates your work to a behavioral ecologist who works on sexual selection in birds, a paleo-ecologist examining climatic regulation of forests, and an ecosystem ecologist who studies the how a dead leaf is transformed into CO2 and minerals? A really cool question that is pitched primarily to mammalogists, mycologists, or myrmecologists, will likely not score particularly high.

B. Broad Concept? Is the question(s) well motivated and justified within a broader conceptual framework?

You may notice that Criteria 1 and 2 share the notion of General and Broad. That’s not a coincidence: what you do must appeal to a wide audience in part, by using the concepts (theory and hypotheses) that structure the field. Once you have posed your question, lay out a suite of hypotheses–complementary and/or contrasting–that outline out your grand plan to deconstruct the question, structure the data you collect, and provide a way of interpreting your results.

Hypotheses are scenarios: statements of a possible reality. The guts of an hypothesis are its assumptions, joined by logic, that yield predictions. You can test hypotheses by testing their assumptions and predictions.

Hypotheses are not a bothersome technicality. They show the PI’s ability to break down the problem into chunks. They have the handy feature of motivating and structuring your data collection.

It is common for a great “A” to ignore or skim through “B”. One result is that the reviewer gets lost in the “Proposed Work” section.

C. Feasible? Does the experimental design/approach logically and feasibly address the question(s) posed?

This is a difficult section to write well, and one of the main reasons why we still have 15-page full proposals. Luckily, the pre-proposal needs just a good first draft of the proposed work, leaving out details of experimental design, time lines, sample sizes, power analyses, etc. That said, the Proposed Work section has to be clear. A common problem in grant proposals also clouds many Materials and Methods sections–details of the work are laid out but without reference to how they answer components of the question and the hypotheses to be tested. The Proposed Work is a good place to brush up on your topic sentences (i.e., To test Hypotheses 2, we will….).

One way to show that something is feasible is to show that you’ve done something like it before. Another is to have pilot data. Pilot data suggests you just didn’t whip this proposal out over Christmas break. The more ambitious the project (and “ambitious project” is a phrase often associated with scores of “VG” and “E”), the greater the need for pilot data.

D. Competent? Are the senior personnel qualified to conduct this research?

Make sure your NSF Bio highlights your expertise and that of your collaborators. Refer a couple of times to this work in the Project Description.

E. Broader Impacts: Benefit society or advance desired societal outcomes

This is still a hard one for me to get a handle on. It is rare for a weak Broader Impacts to sink a great Intellectual Merit. It is not uncommon for a fantastic Broader Impacts to nudge a borderline Do Not Invite into Invite.

A few specifics go a long way. This is a great place to use the Project Summary to  outline what you will do, and the Project Description to give some specifics.

Avoid being vague. It takes as much space to say “I have mentored undergraduates (including 5 females and 5 from underrepresented groups)” as it does to say, “My lab has a long tradition of mentoring females and underrepresented groups.”

If a broader impact can help resource managers or policy makers, contrast briefly how differing results would yield to different advice (in other words, say why this will be helpful). If you are developing K-12 curricula, having a working relationship with your local school district is a big plus.

A few closing thoughts

Program Officers are your allies

Program officers are like Statistics Experts–they’d much rather talk to you as you craft your pre-proposal than help you perform the post-mortem. If you are in the DC area, make an appointment to visit. Volunteer for panel service.

Chasing deadlines

A good pre-proposal needs time to gestate. It will likely benefit from your carving out some writing and thinking time on a monthly basis, rather than investing that same total amount of time to the two or three weeks of Xmas break.

The importance of getting some eyes on your pre-proposal

As the mid-January DEB deadlines approach, you and a lot of your colleagues are in the same boat. You’ve worked hard, crafting each section of your proposal. But with only a week to go, you and only you (and your co-PIs) have likely seen the product. You need to fix that. Here’s one way that works. It is especially germane for senior colleagues in your department who are on the record that “they don’t want pre-proposals to penalize junior colleagues” and are looking for some tangible way to help.

Here’s what we do. A week before the NSF deadline, we assemble a list of department colleagues that are submitting a pre-proposal. We set one day as Review Day. The bargain is that anyone agreeing to review 2-3 proposals–one read-through, about an hour each–gets 2-3 other sets of eyes on their proposal in exchange . The goal here is to identify the big stuff–jargon, poor word choice, “red flags”–that can easily turn a “VG” into a “G” (while you’re at it, market test your title!). The beauty of this system is that the diversity of colleagues in your department is a feature, not a bug, as it comes very close to the diversity of expertise you’d find on a typical NSF panel. I guarantee our “24 hour buddy” system will give you lots of ways, small and large, to craft a better proposal.

Your mileage may vary…

I have been fortunate to serve on a number of NSF panels in my career, and have always found them intensely rewarding experiences. As I said at the beginning, my observations and advice makes sense to me, but should be taken with a grain of salt. I hope this essay catalyzes a conversation. We need that conversation. For, in the end, what makes a quality proposal, like the apocryphal elephant, must to some extent be in the eye of the beholder.

elephant and scientists by Debby Kaspari

Three reviewers trying to understand a very Broader Impact indeed. By Debby Kaspari.

Good luck, and happy grant writing.



A manifesto for Biogeochemical Ecology

The theoretical and empirical challenges of Biogeochemical Ecology include: making biogeochemical maps scalable; translating elemental chemistry into ecological niches; building a science that predicts how the abiotic forms a template for the biotic.  

Biogeochemical Ecology

Twenty-five chemical elements form the recipe for virtually all life on earth; shortfall in any impedes performance and fitness. Gradients of biogeochemistry, combined with those of temperature and precipitation, must therefore create a map of the performance of individuals, the fitness of populations, the structure and function of ecological communities, and the efficacy of ecosystem services. Global Change, in turn, manifests as a massive, continent-scale redistribution and manipulation of temperature, precipitation, and biogeochemistry. My lab’s work focuses on building an empirical and theoretical framework for understanding how Biogeochemical Ecology allows us to predict the consequences of Global Change. This approach provides three basic, interrelated opportunities for transformative science.

  1. Collecting and scaling maps of biogeochemistry according to ecological process. NOAA’s maps of aerosol deposition, and USGS soil maps reveal how natural and anthropogenic drivers rearrange elemental availability on a continental scale that underlie patterns of eutrophication, carbon cycling, and acidification. Finer scale soil studies on the 25 and 50 ha study plots of the Center for Tropical Forest Science reveal the niche structuring of tree communities and their effects on soil biodiversity. Fertilizing still smaller plots from 1600 to 0.25 m2 reveal how the microbes and invertebrates of the brown food web respond in ways that sum to shape decomposition. Our lab pioneered work at each scale, yet scalable maps that link these processes and populations do not yet exist; particularly down to the millimeters and microns relevant to key players like nematodes, fungi, and bacteria. Scalability is one way of recognizing a mature science.
  2. Building and testing a theory of biochemistry-based niches. Despite readily available technology, ecologists know more about the genomes of species, than the elemental composition of species. Yet the niches of organisms are profoundly shaped by their biochemistry: from their investments in P-rich ribosomes and metal-rich enzymes, to chemically distinctive amino acids, to the Ca and P-rich endoskeletons of vertebrates, to the leaky and temperature sensitive Na/K pumps that drive osmoregulation. Our lab has continues to explore the population and ecosystem consequences of two such biochemistry-based niches. One project explores the geographical consequences arising from the Na-poor diet confronting plant consumers. The second project explores the 20-year changes of invertebrate communities across North America as a function of their thermal tolerance, which is enhanced by loading up on P. A third project explores the trophic implications of bioaccumulation: that predators are more likely than their prey to achieve their quota of metals like Na, Cu, and Zn. All combine metabolic approaches that search for commonalities rooted in deep history of plants and animals, but with a natural historian’s eye for detail. When combined with scalable maps of biogeochemistry, they hold the potential for a deep understanding of the geographical ecology of abundance and ecosystem function.
  3. Fusing the three great classes of abiotic drivers into a useable theory predicting the synergy of multiple stressors. A hallmark of global change is the recombination of abiotic drivers to form new environments. Perhaps our greatest challenge is understanding how phenotypic plasticity, ecological filters, and natural selection shape populations and ecosystems particularly when we move beyond a singular focus on one driver like temperature or drought. In short, when do multiple stressors act like drug cocktails to hinder adaptive responses of organisms? We seek to understand the degree to which thermal and drought tolerance can be built from biochemical parts and how often-hidden patterns of malnutrition ramify to decrease resilience in populations and ecosystems.

To paraphrase GE Hutchinson, one of the great challenges for modern ecology is a deep understanding of how the Ecological Play is constrained by the Biogeochemical Theater.

Know Your Grassland Invertebrates: The Cobweb Spiders

An occasional series by AntLab Parataxonomist: Brittany Benson

Theridiidae_BlackMask_SaltGrassJuly2015_F_D_110556 Theridiidae_BlackMask_SaltGrassJuly2015_V_110556

Theridiidae – Cobweb Spiders, or Comb-Footed Spiders

Have a comb of serrated setae on the fourth tibiae (these are used to draw out and fling silk when attacking).  Also, typically have a round opisthosoma, but some have a tear-drop shape or weird bumps along with a triangle shape.

32 genera in North America with over 200 species.  One of the most diverse spider families worldwide, this family has more than 2,200 species.  The common house spider is in this family (this spider lives with humans on every continent save for Antarctica, and is possibly the most common spider in the world).  Though most are harmless, it is important to note that this also the family that the widows are members of – these being in the genus Lactrodectus.  Some theridiids are kleptoparasites in the webs of other spiders, eating their prey and sometimes the host spider.  The three-dimensional aspect of theridiids may seem chaotic, but is actually quite fantastic – prey gets stuck to a strand, and its struggling breaks the strand at a weak point causing it to be pulled up to the next strand, now suspended in the web (if the prey got stuck by trying to walk under the web), and the spider rushes in to wrap it up.  They love to make their homes in sheltered areas, and, back when outhouses were still the thing, one normally took a stick along to remove any webbing before sitting –  this seemed to be a great hangout for widows.  I was told the crackling sound the web made as you brushed a stick through it let you know you had a widow, but I now wonder if that could be for all theridiids – either way, better safe than sorry, especially when it’s the middle of the night, and you had to walk a ways into the trees to even get to the outhouse.  Hooray for indoor plumbing!

However, the cobweb architecture that comes to mind is not the only web form in this family – there is actually quite a bit of diversity in web type, which is unusual for within a family.  It is also interesting to note that there are even permanently social spiders in the Theridiidae.  

Know Your Grassland Invertebrates: The Checkered Beetles


An occasional series by AntLab Parataxonomist: Brittany BensonCleridae_SaltGrassJuly2015_NormanOK_D_110409Cleridae_SaltGrass2015_NormanOK_V_110409

Cleridae – Checkered Beetles

With more than 3,500 species in 330 genera worldwide, about 300 species in 36 genera occur in North America, being more diverse in the south.  They have a vast size range of 2-24mm, but overall have a distinct, elongated body type (even th more robust ones are still distinctly clerid-shaped).  They are also covered with hairs, and can be very  neatly patterned and colored (thus the name of checkered beetles).  

Most of the members of this family are predacious on other insects, both as larvae and adults.  Typically found with woody things (check out the surface and under bark for an easy sighting).  Just about any part of the woody plant they can get to, they’ll be there.  Have a pile of dead twigs?  There may be a clerid there.  Some are fierce predators of bark beetles, and so may be important in controlling those populations.  Adults of some (mostly in the Clerinae) prefer to live on flowers and munch on the pollen.  Even if the larvae and adults are both typically predacious, they may not feed on the same things.  One species of Aulicus, for example, eats the eggs of lubber grasshoppers as larvae, and the adults take to eating noctuid caterpillars.  Some species are even scavengers.

Know Your Grassland Invertebrate: The Leafhoppers

An occasional series by AntLab Parataxonomist: Brittany Benson

Cicadellidae_SaltGrass2015_NormanOK_110522_D Cicadellidae_SpeckledSand_SaltGrass2015_NormanOK_110522_V


Hind tibiae with 1 or more rows of small spines – this is the main thing separating this family from the other Cicadoidea families.

Commonly called leafhoppers, there are about 3,000 North American species, around 22,000 worldwide.  They can vary in size from 2mm to 30mm, but most stay under 13mm.  These are hemimetabolous insects, so their nymphs look like miniature versions of the adults, but with different proportions (usually larger heads), and the wings have not developed yet – again, do not mistake the developing wing pads for wings.  Wings will be as long or longer than the body, where the wing pads leave much or most of the abdomen exposed.

Occurring on nearly all types of plants – trees, shrubs, grasses, fields, gardens – the leafhoppers can be found in about every habitat that has vascular plants, as varied as deserts and wetlands.

They feed mainly on the leaves of their host plant, most species being quite specific, so habitat for a species can also be well defined.  Many act as vectors for plant diseases.

  Plenty of them emit honeydew from their anus.  Yummy!  It’s a cocktail made from unused plant sap and the insect’s own special waste products.  Surprisingly, there are few known instances of ants tending these fru fru drink specialists.  One Brazilian species was observed to be tended by up to 21 different ant species, but it seems as though it is an opportunistic relationship.  But another example, that of Dalbulus quinquenotatus, shows that some are complete myrmecophiles – this species will allow ants to approach, and when the ants stroke them with their antennae, the leafhopper nymphs will secrete a honeydew droplet and hold it in place until an ant takes it.  Nonattended species of Dalbulus will avoid ants by walking away, hopping, or jumping.  The droplets of the ant-loving species are also much larger and more frequently excreted than in other species.  

Many can produce sound, but they are weak sounds (at least to us, and we should totally judge the strength of other and utterly different creatures to that of our own).  These sounds are produced by vibrating timbals on the dorsolateral base of the abdomen.  Tymbals are thin-walled bits of the body wall.  They can be sounds of disturbance, courtship, and lady sounds, but all the sounds appear to be species specific.  

Two More New Positions in Geographical Ecology

Our friends in the Department of Geography and Environmental Sustainability continue to grow, and have posted two hires out of the 19 total in Environmental Science at OU (including Biology’s own two in Geographical Ecology, Deadline 1 October!).  Join us as we build something great!

Tenure-track Assistant or Associate Professor in Geohumanities and Tenure-track Assistant or Associate Professor in Socio-Ecological Systems Modeling

OKLAHOMA, NORMAN 73019. The Department of Geography and Environmental Sustainability at the University of Oklahoma ( is an actively growing and dynamic department. Over the past 5 years, undergraduate enrollment has more than tripled, research expenditures have doubled and the department has quintupled its computer resources for research and teaching.  As part of our ongoing growth, applications are being solicited for two tenure-track assistant or associate professors.


Position 1: Tenure-track Assistant or Associate Professor in Geohumanities

We seek a geohumanities scholar who studies environment-society relations, utilizes critical theory, and incorporates stakeholder/community engagement. An urban or regional specialization would be welcome. The successful candidate will have an opportunity to participate in new and established campus-wide efforts to support humanities scholarship. To raise our research profile, we desire faculty members who are strong in research in terms of suitable publications and grant success, as well as active within both professional and stakeholder communities and effective at bringing research insights into the classroom.

Teaching duties would include introductory geohumanities courses as well as a course on environment and society, and possibly regional courses within the candidate’s expertise.

Application Process: Candidates are invited to submit a statement of interest and qualifications, a full curriculum vita, up to five scholarly publications, and a list of three references. Screening will begin September 15th, 2015. To apply, please submit applications electronically in one PDF file to Dr. Laurel Smith, Chair, Human Geography Search Committee at Please copy Ms. Deborah Marsh at


Position 2: Tenure-track Assistant or Associate Professor in Socio-Ecological Systems Modeling

The successful candidate will be capable of research excellence in socio-ecological systems modeling as related to climate variability and change, and they must have demonstrated success in proactively leading and/or participating in interdisciplinary, collaborative teams. In particular, the candidate will fill an immediate need in coupled human-natural systems modeling, preferably through Bayesian hierarchical modeling or similar methodology.  The candidate will also be able to work within collaborative teams that examine challenging questions facing the social, natural, and physical sciences.

Application Process: Candidates are invited to submit a statement of interest and qualifications, a full curriculum vita, copies of up to five scholarly publications, and a list of three references. To apply, please submit all materials electronically in one PDF file to Dr. Bruce Hoagland, SES Search Committee Chair (Email: and copy Ms. Deborah Marsh (

Screening will begin October 1st, 2015. Applicants for both positions should have demonstrated a commitment to teaching at the undergraduate and graduate levels and a willingness to participate in Department, University, and professional service. We seek an outstanding candidate with a PhD in geography, or closely related field.

Persons from under-represented groups are strongly encouraged to apply. Initial appointment to this position will begin August 2016. Salary and remunerations are competitive and commensurate with qualifications.


Candidates interested in collaborative research will find many exciting opportunities on the campus. The University hosts the History of Science Collections, Digital Scholarship Lab in the Bizzell Memorial Library, Western History Collections, Carl Albert Congressional Research and Studies Center, and the Fred Jones Jr. Museum of Art. The College of Atmospheric and Geographic Sciences hosts the National Weather Center, South Central Climate Science Center, Southern Climate Impacts Planning Program, and the Center for Spatial Analysis. The Department is home to the Oklahoma Wind Power Initiative and the Land Use Land Cover Institute.


The University of Oklahoma (OU) is a Carnegie-R1 comprehensive public research university known for excellence in teaching, research, and community engagement, serving the educational, cultural, economic and health-care needs of the state, region, and nation from three campuses: Norman, Health Sciences Center in Oklahoma City and the Schusterman Center in Tulsa. OU enrolls over 30,000 students and has more than 2700 full-time faculty members in 21 colleges. In 2014, OU became the first public institution ever to rank #1 nationally in the recruitment of National Merit Scholars, with 311 scholars. The 277-acre Research Campus in Norman was named the No.1 research campus in the nation by the Association of Research Parks in 2013. Norman is a culturally rich and vibrant town of around 113,000 inhabitants located just outside Oklahoma City.  With outstanding schools, amenities, and a low cost of living, Norman is a perennial contender on “best place to live” rankings. Visit and for more information.

The University of Oklahoma is an equal opportunity institution


10 Foundational Papers in Community Ecology

I suggest to students in Advanced EEB, our course for first semester Ph. D. students, that creativity is about fostering

  1. your ability to generate associations/ideas, and
  2. your judgement as to which ideas are worth pursuing.

Toward that end, I also encouraged them to ask professors for a list of ten influential papers (“papers everybody should read” is how I think I put it).

Then, of course, BeccaP immediately sends me an email asking for my list. Serves me right.

The following ten papers occurred to me on a Saturday morning, over 10 minutes or so, and in the middle of my third cup of coffee. I figured they must be important to me if they sprung up with no more prompting than a dose of caffeine. They are not my “Top 10” (an odd concept), but they certainly exist in my 99th percentile.

Turns out, these kinds of lists serve both elements of the creative enterprise. First, the papers below are all full of ideas (some ultimately more successful than others) and represent highly creative people at the height of the powers. I remember, or at least I think I do, reading each for the first time and feeling energized and a little bit jealous.  At the same time, they reflect my judgement as to the elements that combine to form a good science paper: clarity, ambition, stepping boldly into a knowledge gap, identifying and advocating a path forward. Each photocopy was highly scribbled upon. You can clearly see the fingerprints of these ecologists all over our lab’s work.  The first on the list made such an impression,  I deconstructed it.

MacArthur, R. H. 1958. Population ecology of some warblers of northeastern coniferous forests. Ecology 39:599-619.

Hutchinson, G. 1959. Homage to Santa Rosalia, or why are there so many kinds of animals? American Naturalist 93:145-159.

Hairston, N., F. Smith, and L. Slobodkin. 1960. Community structure, population control, and competition. American Naturalist 94:421-425.

Janzen, D. 1967. Why mountain passes are higher in the Tropics. American Naturalist 101:233-249.

McNaughton, S., M. Oesterheld, D. Frank, and K. Williams. 1989. Ecosystem-level patterns of primary productivity and herbivory in terrestrial habitats. Nature 341:142-144.

Power, M. E. 1992. Top-down and bottom-up forces in food webs: do plants have primacy? Ecology 73:733-746.

Holling, C. 1992. Cross-Scale Morphology, Geometry, and Dynamics of Ecosystems. Ecological Monographs 62:447-502.

Ritchie, M. E. and H. Olff. 1999. Spatial scaling laws yield a synthetic theory of biodiversity. Nature 400:557-560.

McGill, B. J., B. J. Enquist, E. Weiher, and M. Westoby. 2006. Rebuilding community ecology using functional traits. Trends in Ecology and Evolution 21:175-185.

Orians, G. H. and A. V. Milewski. 2007. Ecology of Australia: the effects of nutrient-poor soils and intense fires. Biological Review 82:393-423.

The tropics is just a big plate of scrambled eggs

Debby Kaspari is a featured artist at the Leigh Yawkey Woodson’s annual “Birds in Art” competition. Here is a brief video that captures the glories of painting birds in tropical forests.