For the past several years we have been investigating effects of prescribed fire on native ground-nesting bees in remnant prairies and restorations across our fragmented prairie landscape. We have a paper about effects of fire on the abundance and diversity of nesting bees in prep for a peer-reviewed journal. We are posting recommendations from this investigation now as a one-page non-technical document.
Funding for this project was provided by the Minnesota Environment and Natural Resources Trust Fund as recommended by the Legislative-Citizen Commission on Minnesota Resources (LCCMR) and by the National Science Foundation.
For the past few years we have been studying effects of prescribed fire on pollination, pollinators, and plants across our fragmented prairie landscape. We have a paper about effects of fire on pollen that we are preparing to submit to a peer-reviewed journal. We are posting recommendations from this investigation now as a one-page non-technical document.
Funding for this project was provided by the Minnesota Environment and Natural Resources Trust Fund as recommended by the Legislative-Citizen Commission on Minnesota Resources (LCCMR) and by the National Science Foundation.
For the past few years we have been studying effects of prescribed fire on pollination, pollinators, and plants across our fragmented prairie landscape. We have three papers we are preparing for submission to peer-reviewed journals. From each, we have recommendations for land managers interested in conserving native prairie bees and plants. We are posting these recommendations now in a one-page non-technical format.
The first paper is about prescribed fire effects on pollination and pollinator visitation. Read the recommendations that are based on the paper.
Expect two more one-pagers tomorrow.
Funding for this project was provided by the Minnesota Environment and Natural Resources Trust Fund as recommended by the Legislative-Citizen Commission on Minnesota Resources (LCCMR) and by the National Science Foundation.
Last week I trekked out to Landfill to see if I could acquire some aphids and I was successful! The most successful way of harvesting them in the past which was using a paintbrush that had a small amount of bristles. However, I found most successful for preparing the Aphids to move is slightly blowing on them. This provoked them enough to retract their styles so I could sweep them using a larger paintbrush. After a drive back, I put them back onto the leaves of the addition plants in ExPt01.
This week I was met with some pretty unfortunate news after visiting the addition plants in ExPt01, they were no longer present. So to remedy this, I will venture back out to landfill and harvest more aphids and check back in with them tomorrow afternoon to make sure that they are thriving on their respective plants.
The Echinacea Project, and its principal investigator Stuart Wagenius, is responsible for the foundation of one of the most comprehensive and long-running studies of prairie plants. Experimental plots use a common garden design and include experiments involving inbreeding, aphid addition and exclusion, flowering phenology, pollen addition and exclusion, and more. The Echinacea angustifolia in experimental plot 1 were planted as far back as 1996; while they have experienced mortality over the years, many of these plants are still alive. Team Echinacea, most of which are younger than the plants we work with, have been hard at work collecting measurements to add to the ever-expanding dataset. We take data on things like the number of rosettes, leaves, and flowering heads a plant has, as well as insect activity and disease spread, to help determine the fitness of the plants in various treatments.
A common wood-nymph being thwarted by our pollinator-exclusion bagsAn Agapostemon virescens visiting echinacea in ExPt01A fungus commonly found growing in ExPt01A jumping spider on the hunt, likely preying on floral visitorsDragonflies, abundant in ExPt08Aaron, having a standoff with a grasshopper
As of August 1, 2025, we have measured approximately 60% of experimental plot 1, which has over 11,000 planted positions. Having accomplished so much, we plan on beginning harvesting soon next week. After the heads are harvested, the achenes will be sent to the Chicago Botanic garden, where interns and volunteers will work to determine the flowers’ seed set. Throughout the rest of the field season, Team Echinacea will continue measuring in ExPt01, and begin on ExPt08, ExPt09, ExPt07, and ExPt02.
This is a map of measuring progress in ExPt01, where the blue segments have been completed and the white, unfilled sections are soon to be measured. We measure by experiment, concluding one before we move to another. We have completed the inbreeding 1 cohort, the inbreeding 2 cohort, and the first generation of the heritability of fitness experiment. Currently, we are working on the 99 main garden, a series of rows planted by Stuart in 1999.
The aphid addition and exclusion experiment was started in 2011 by Katherine Muller. The original experiment included 100 plants selected from exPt01 that were each assigned to have aphids either added or excluded across multiple years. The intention is to assess the impact of the specialist herbivore Aphis Echinaceae on Echinacea fitness.
In 2025, 32 of the original 100 plants were alive, 12 addition and 20 exclusion plants. Unfortunately, no aphids were found in exP01 but many aphids were found in other remnant plots this year. As soon as I can I will transfer aphids from the plants out in remnants to plants in exP01 to get this experiment on the move.
Plant status (basal, flowering, not present), aphids present, ants present, herbivory (number of leaves significantly chewed on), and the number of aphids added/removed (depending on specific treatment)
Protocols and datasheets are located at ~Dropbox\aphidAddEx\aphids2025
Samples collected: NA
Products:
Andy Hoyt’s poster presented at the Fall 2018 Research Symposium at Carleton College
2016 paper by Katherine Muller and Stuart on aphids and foliar herbivory damage on Echinacea
2015 paper by Ruth Shaw and Stuart on fitness and demographic consequences of aphid loads
Team Echinacea has begun the annual census of Echinacea plants. Each year we census all flowering plants at over 30 prairie sites. Each plant gets a digital census record, a flag, and a tag. Most plants already have a tag, so we don’t give them a new one. But some have lost their tag or are flowering for the first time, so they need a new tag. New tags this year are numbered starting at 30,001. Plants get neon flags and we will come back and survey them so we can make a map of the location of every plant. Once they get surveyed, we replace the neon flag. All of these efforts help build a long-term dataset about the survival and reproduction of these very long-lived plants. These plants face many challenges living in small prairie patches, but they are tough.
Below is a summary of the number of census records taken so far at nine sites
site rawSite demo.id
1 other 1
2 alf around landfill 11
3 cg common garden 23
4 eelr east elk lake road 19
5 lfe landfill east 118
6 lfw landfill west 99
7 lce loeffler corner east 78
8 lcw loeffler corner west 81
9 rrxx railroad crossing 36
Amy Waananen’s just had a paper published in New Phytologist. She reports how the difference in flowering time of a plant’s two parents influences the plant’s fitness. Many researchers have investigated how differences in the location of parents (close or far) influences progeny fitness. Few, if any, have investigated how differences in flowering time affects progeny fitness.
Differences in flowering time are not as straightforward as differences in location. In space, parents can be far or close and we use distance to measure it. To state the obvious, negative distances do not makes sense. In time, parents can be far (asynchronous) or near (synchronous). But maternal plants can be earlier or later than paternal plants. This aspect of distance in time is fundamentally different than distance in space. Amy used positive and negative values to indicate which parent flowered earlier. Remarkably, this perspective really mattered to the fitness of progeny. Wow!
Amy’s discovery is really cool, it’s a surprise, and it’s a useful contribution to basic science. Amy also suggested some non-intuitive management strategies that can help promote plant fitness and resilience of populations in the face of changing environments.
Here’s the citation:
Waananen, A., Ison, J.L., Wagenius, S. and Shaw, R.G. 2025. The fitness effects of outcrossing distance depend on parental flowering phenology in fragmented populations of a tallgrass prairie forb. New Phytologist. https://doi.org/10.1111/nph.70240
A rainbow-like cloud glows over exPt01 while the team was in MN for fall 2023 burns. A good omen for the field season that followed!
Introduction
Every year since 1996, members of Team Echinacea have recorded flowering phenology, taking measure data and harvested heads from thousands of Echinacea angustifolia plants in plots with “common garden” experimental designs. These experimental plots are located in prairie remnants, restorations, and abandoned agricultural fields that are managed as grassland habitat. Currently, the Echinacea Project has 10 established experimental plots. Some plots have multiple ongoing experiments within.
In the past few years, we have scaled back significantly on taking phenology records in the experimental plots. During 2023 and 2024, our primary goals with phenology were to a) map out the positions of flowering plants/heads within the plots, b) deploy twist ties to all flowering heads to ease measuring and harvesting, and c) to record the day of first flowering for all heads in plots to continue the long-term data collection in a more scaled-back fashion. As a result, we conducted only a few rounds of phenology per plot and did not capture the full range of flowering dates for every head. Phenology info is briefly reported on in each plot’s update along with the location of the data. This applies to the hybrid experimental plots as well.
Crew members Wyatt and Emma search can’t find positions in exPt01
Experimental plot 1 was first planted in 1996 (cleverly termed the 1996 cohort), and has been planted with nine other experiments in subsequent years, with the most recent planting being Amy Waananen’s inter-remnant crosses. It is the largest of the experimental plots, with over 10,000 planted positions; experiments in the plot include testing fitness differences between remnants (1996, 1997, 1999), quantifying effects of inbreeding (inb1, inb2), and assessing quantitative genetic variation (qGen1). It also houses a number of smaller experiments, including fitness of Hesperostipa spartea, aphid addition and exclusion, and pollen addition and exclusion (these experiments have separate posts).
In 2024, we conducted phenology in this plot between July 8th and July 18th. During measure, we visited 3123 of the 10,992 positions planted and found 2728 living plants. 83 plants were classified as “flowering” in exPt01 this year, totally 96 heads. This is a significantly fewer plants than flowered in summer 2023 (560). In summer 2024, we harvested 82 total Echinacea heads in exPt01 (including many from the ever-productive 99 south garden).
ExPt01 is also the only plot to have staples marking positions where plants that have died used to be. We added 67 staples to the experimental plot this year, but only in locations that we couldn’t find staples during measure that were already supposed to be there. We didn’t have time to get to every position with a missing staple (see where we covered here: Dropbox/CGData/125_measure/measure2024/staple2024/2024addStaplesExPt01.pdf). We did not have time to re-search locations that we called plants “can’t finds” at three years in a row in 2023 and 2024. Once these locations receive their final search, hopefully in 2025, we can put staples at them as well.
This experiment was started in 2020 by Amy Waananen to understand how the distance between plants in space and in their timing of flowering influences the fitness of their offspring. If plants that are located close together or flower at the same time are closely related, their offspring might be more closely related and inbred, and have lower fitness than plants that are far apart and/or flower more asynchronously. Plants in this experiment resulted from interremnant hand-crossings from 9 remnants: On27, SGC, GC, NGC, EELR, KJ, NNWLF, NWLF, LF. Crossing took place in 2020 and 2021, and individuals were planted in 2020 (as seed) and 2022 (as plugs). Surviving plants were assigned cgPlaIds in 2023 and incorporated into the p1 workflow. Mortality in this experiment has been high, with 80.3% of positions searched in 2024 resulting in “can’t finds.”. We did not re-search “can’t find” positions in 2024 due to time constraints (with Amy’s approval).
The inb1 experiment investigates the relationship between inbreeding level and fitness in Echinacea angustifolia. Each plant in experiment inb1 originates from one of three cross types, depending on the relatedness of the parents: between maternal half siblings; between plants from the same remnant, but not sharing a maternal or paternal parent; and between individuals from different remnants. All individuals were planted in 2001. We continued to measure fitness and flowering phenology in these plants.
The inb2 experiment investigates the relationship between inbreeding level and fitness in Echinacea angustifolia. Each plant in experiment inb2 originates from one of three cross types, depending on the relatedness of the parents: between maternal half siblings; between plants from the same remnant, but not sharing a maternal or paternal parent; and between individuals from different remnants. All individuals were planted in 2006. We continued to measure fitness and flowering phenology in these plants. In October 2024, former team member Riley Thoen recently published a paper in the Journal of Hereditary on the conservation value of small remnants using results from this experiment.
The qGen1 (quantitative genetics, or just qGen) experiment in p1 was designed to quantify the heritability of traits in Echinacea angustifolia. We are especially interested in Darwinian fitness. Could fitness be heritable? During the summer of 2002 we crossed plants from the 1996 & 1997 cohorts of exPt01. We harvested heads, dissected achenes, and germinated seeds over the winter. In the spring of 2003 we planted the resulting 4468 seedlings (this great number gave rise to this experiment’s nickname “big batch”).
data in cgData repo: ~/cgData/summer2024/exPt01Phenology
Measure data (status, size, etc.)
data in SQL database
Harvest data (IDs of harvested heads, missing achenes, etc)
detailed data in dropbox: dropbox/CGData/140_reconcile/reconcile2024/reconcileOut/2024harvestListReconciledExport.csv
data in SQL database
data in echinaceaLab package (hh.2024)
Samples collected:
82 heads harvested
At cbg for processing (counted, ready to randomize)
Products:
Publications
Thoen, R. D., A. Southgate, G. Kiefer, R.G. Shaw, S. Wagenius, The conservation value of small population remnants: Variability in inbreeding depression and heterosis of a perennial herb, the narrow-leaved purple coneflower (Echinacea angustifolia). 2024. Journal of Heredity esae055. https://doi.org/10.1093/jhered/esae055.
Page, M. L., Ison, J. L., Bewley, A. L., Holsinger, K. M., Kaul, A. D., Koch, K. E., Kolis, K. M., and Wagenius, S. 2019. Pollinator effectiveness in a composite: A specialist bee pollinates more florets but does not move pollen farther than other visitors. American Journal of Botany 106: 1487–1498. PDF
Waananen, A., G. Kiefer, J. L. Ison, and S. Wagenius. 2018. Mating opportunity increases with synchrony of flowering among years more than synchrony within years in a nonmasting perennial. The American Naturalist 192: 379-388. PDF | Appendix | online version
Muller, K. and S. Wagenius. 2016. Echinacea angustifolia and its specialist ant-tended aphid: a multi-year study of manipulated and naturally-occurring aphid infestation. Ecological Entomology 41: 51-60. PDF | online version
Shaw, R. G., S. Wagenius and C. J. Geyer. 2015. The susceptibility of Echinacea angustifolia to a specialist aphid: eco-evolutionary perspective on genotypic variation and demographic consequences. Journal of Ecology 103: 809-818. PDF
Kittelson, P., S. Wagenius, R. Nielsen, S. Qazi, M. Howe, G. Kiefer, and R. G. Shaw. 2015. Leaf functional traits, herbivory, and genetic diversity in Echinacea: Implications for fragmented populations. Ecology 96: 1877–1886. PDF
Ison, J.L., and S. Wagenius. 2014. Both flowering time and spatial isolation affect reproduction in Echinacea angustifolia. Journal of Ecology 102: 920–929. PDF | Supplemental Material | Archived Data
Ison, J.L., S. Wagenius, D. Reitz., M.V. Ashley. 2014. Mating between Echinacea angustifolia (Asteraceae) individuals increases with their flowering synchrony and spatial proximity. American Journal of Botany 101: 180-189. PDF
Ridley CE, Hangelbroek HH, Wagenius S, Stanton-Geddes J, Shaw RG, 2011. The effect of plant inbreeding and stoichiometry on interactions with herbivores in nature: Echinacea angustifolia and its specialist aphid. PLoS ONE 6(9): e24762. http://dx.plos.org/10.1371/journal.pone.0024762
Wagenius, S., H. H. Hangelbroek, C. E. Ridley, and R. G. Shaw. 2010. Biparental inbreeding and interremnant mating in a perennial prairie plant: fitness consequences for progeny in their first eight years. Evolution 64: 761-771. Abstract | PDF
Ruth G. Shaw, Charles J. Geyer, Stuart Wagenius, Helen H. Hangelbroek, and Julie R. Etterson. 2008. Unifying life-history analyses for inference of fitness and population growth. American Naturalist 172: E35 – E47. Abstract | PDF | Supplemental Material
Geyer, C.J., S. Wagenius, and R.G. Shaw. 2007. Aster models for life history analysis. Biometrika 94: 415-426. PDF | Supplemental Material
Grad student work
Drake Mullett’s PhD dissertation (2025)
Wyatt Mosiman’s MS thesis (2024)
Amy Waananen’s paper “The fitness effects of outcrossing distance depend on parental flowering phenology in fragmented populations of a tallgrass prairie forb” (with co-authors Ison, Wagenius, and Shaw) was just accepted by New Phytologist–it includes data from parents in exPt01 and progeny in exPt02.
AKA the heritability of flowering time experiment, exPt02 was designed to examine the role flowering phenology plays in the reproduction of Echinacea angustifolia. Jennifer Ison planted this plot in 2006 with 3,961 individuals selected for extreme (early or late) flowering timing, known as phenology. Using this phenological data, we explore how flowering phenology influences reproductive fitness and estimate the heritability of flowering time in E. angustifolia. In the summer of 2024, we conducted phenology between July 10th and July 19th. During measure, we visited 1,725 positions of the 3,961 positions originally planted. We measured 1,190 living plants, of which 302 were flowering with a total of 402 flowering heads (count excludes vertical developments). In the fall, we harvested 375 heads from exPt02. We observed much lower levels of seed predation by ground squirrels this year than the past few years.
data in cgData repo: ~/cgData/summer2024/exPt02Phenology
Measure data (status, size, etc.)
data in SQL database
Harvest data (IDs of harvested heads, missing achenes, etc)
detailed data in dropbox: dropbox/CGData/140_reconcile/reconcile2024/reconcileOut/2024harvestListReconciledExport.csv
data in SQL database
data in echinaceaLab package (hh.2024)
Samples collected:
375 heads harvested
at CBG for processing
Products:
Papers
Pearson, A.E., Z. Zelman, L.A. Hill, M.A. Stevens, E.X. Jackson, M.M.N. Incarnato, R.M. Johnson, S. Wagenius, and J.L. Ison. 2023. Pollinators differ in their contribution to the male fitness of a self-incompatible composite. American Journal of Botany 110(6): e16190. https://doi.org/10.1002/ajb2.16190
Reed, W. J., J. L. Ison, A. Waananen, F. H. Shaw, S. Wagenius, R. G. Shaw. 2022. Genetic variation in reproductive timing in a long-lived herbaceous perennial. American Journal of Botany 109(11) 1861–1874: https://doi.org/10.1002/ajb2.16072
Page, M. L., Ison, J. L., Bewley, A. L., Holsinger, K. M., Kaul, A. D., Koch, K. E., Kolis, K. M., and Wagenius, S. 2019. Pollinator effectiveness in a composite: A specialist bee pollinates more florets but does not move pollen farther than other visitors. American Journal of Botany 106: 1487–1498. PDF
Grad student work
Wyatt Mosiman’s MS thesis (2024)
Amy Waananen’s paper “The fitness effects of outcrossing distance depend on parental flowering phenology in fragmented populations of a tallgrass prairie forb” (with co-authors Ison, Wagenius, and Shaw) was just accepted by New Phytologist–it includes data from parents in exPt01 and progeny in exPt02.
We may have missed some other products.
exPt05:
The only experimental plot at Staffanson Prairie Preserve (SPP), exPt05 was planted to compare progeny of maternal plants from burned and unburned sections of SPP. There were originally 2800 individuals planted, but high mortality made it impractical to visit the plot row-by-row. Now, we treat the plot like demography. We use our survey-grade GPS to find plants in exPt05 that have previously flowered and add more plants to the stake file if new plants in the plot flower. In 2024, we found 17 living plants in exPt05 during flowering/total demo, 8 of which were flowering! We also observed (but did not take data on) additional basal plants within the plot boundaries that appeared to be growing on a 1×1 meter grid. ExPt05 persists!
Team Echinacea established quantitative genetics experiments to quantify additive genetic variance of fitness in Echinacea, with the idea that we can estimate evolutionary potential of study populations. The plants in qGen2 and qGen3 are plants in the 1996, 1997, and 1999 cohorts. These plants were crossed with pollen from plants in remnants to produce seed for qGen2 and qGen3, which now inhabit exPt08. Originally, 12,813 seeds were sown in the common garden. Seeds from the same cross (shared maternal and paternal plants) were sown in meter-long segments between nails. In the summer of 2024, we conducted phenology between July 10th and July 19th. A total of 3,253 seedlings were originally found, but due to gradual mortality we only searched 448 positions in 2024 for plants, and we found evidence of 314 living individuals. We identified 24 flowering plants in with a total of 29 heads, of which we harvested 23.
data in cgData repo: ~/cgData/summer2024/exPt08Phenology
Measure data (status, size, etc.)
data in SQL database
Harvest data (IDs of harvested heads, missing achenes, etc)
detailed data in dropbox: dropbox/CGData/140_reconcile/reconcile2024/reconcileOut/2024harvestListReconciledExport.csv
data in SQL database
data in echinaceaLab package (hh.2024)
Samples collected:
23 heads harvested
at CBG for processing
Products:
None… yet!
tplot
tplot is located within the bounds of exPt08. Plants of many species here were rescued from the landfill site. They we transferred as chunks of prairie sod, and individual transplants. This year during flowering and total demo, we encountered 14 living Echinacea plants from which we harvest 12 heads.
Demography data: head counts, rosette counts, etc.
demap input files have been updated with 2024
Spatial location for all flowering and some basal (total demo) plants
demap input files have been updated with 2024
Harvest data (IDs of harvested heads, missing achenes, etc)
detailed data in dropbox: dropbox/CGData/140_reconcile/reconcile2024/reconcileOut/2024harvestListReconciledExport.csv
data in echinaceaLab package (hh.2024)
Samples collected:
12 heads harvested
at CBG for processing
Products:
None… yet!
Experimental plot management:
Just like other areas of the prairie, our experimental plots need management! Here’s a list of the stewardship activities that we conducted in or for our plots during 2024:
NU MS student Maddie Sadler shows off our sweet clover haul
None of our experimental plots burned in the fall or spring prior to the 2024 growing season
Collected seed to plant in p1, p2, & p8, including:
1) Elise collected Carex brevior, Carex bicknellii, and Carex gravida from several sites and from plants that we established a few years ago near exPt01
2) Liam established Viola pedatifida production tub,
3) We collected a few additional species (including Galium boreale, Solidago missouriensis, Astragalus adsurgens, Dichanthelium leibergii, and Bromus kalmii) we plan to establish via plug in spring 2025
Psst – next year person writing this report; I ran my numbers using the script Dropbox/echProjAdmin/projectStatusReports/psr2024/wmGatherDataForReports2024.R. Maybe this can help you out.
Team Echinacea has successfully completed some spring prescribed burns! Our mighty team of seven (Stuart, Gretel, Jared, Wyatt, Fannie, Brad, and I) took to Minnesota this past week to conduct a handful of spring prescribed burns. We set off bright and early on Sunday morning at around 9am to make the 9 hour drive from Chicago Botanic Garden to our study sites in Minnesota. Weather in Minnesota on Monday and Tuesday allowed us to get multiple different sites done including the coveted P1 and P8 experimental plots. The sun was shining, the sky was clear, humidity was low, the wind was blowing just enough, and the grass was very dry making it a perfect day. The days were long and the temperatures were high (in the 80’s), but we polished off both days with some great food and great conversations which are essential to any good burn trip to keep up morale.
Experimental plot p1 during the burn (left) and after the burn (right)
Fannie using the drip torch for the first time (left) and Jared lecturing Blue on the importance of fire safety (right)
Not only did we get some good burns in, but we were able to visit Runestone County Park on Tuesday morning. We used this trip to see the current restoration work being done at the park, and it allowed us to find spots for potential signs discussing different topics such as why prescribed fire matters, the history of prairies, and more. We also used our time in MN to get many pictures and videos to be used for dissemination projects discussing why prescribed fire is important for native pollinators.
Of course it wouldn’t be a complete trip to Minnesota without a stop at Staffanson Prairie Preserve.
This is part of our project “How Do Prescribed Fires Affect Native Prairie Bees?”
Funding for this project was provided by the Minnesota Environment and Natural Resources Trust Fund as recommended by the Legislative-Citizen Commission on Minnesota Resources (LCCMR).
