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.
2024 was a relatively low-flowering year in the common gardens, but that doesn’t mean we didn’t bring plenty of Echinacea heads back to the lab for our volunteers! Here are our stats:
TOTAL: 689heads
exPt01: 82 heads (11.9%)
exPt02: 375 heads (54.4%)
exPt07: 87 heads (12.6%)
exPt08: 23 heads (3.3%)
exPt09: 110 heads (16.0%)
tplot: 12 heads (1.7%)
(rounding errors in %s abound)
Our first harvest was at p2, p7, and p9 on August 19th. Our last harvest was all the way on October 3rd at p7 and p9! Those hybrid plants really held on for awhile. Unfortunately, our first harvest was not in time to harvest some heads before they begin dispersing, and there are always a couple that lose achenes throughout the harvest season before we catch them. This year wasn’t toooo egregious, but our worst head lost an estimated 85% of its achenes before we encountered it. Yikes! Keep an eye out for seedlings around (7, 2) in p2…
A few achenes shy of a full head…
One major victory from this year is that, besides a couple holes and piles of dirt, the ground squirrels were relatively well behaved in p2 this year. Typically, they enjoy ripping heads off echinacea plants and leaving stems, heads, and achenes everywhere. Based on harvest notes, in 2024 we only found two heads on the ground and only one was missing entirely. Compare this to 2022, when the team only harvested 41% of the heads they planned on harvesting due to ground squirrel activity. (Though, I suspect this may have been a blessing—they still harvested 480 heads. If they’d harvested all of them, it would have been around 1,170 heads. That’s a lot of harvesting!)
Another fun fact: The 99 south garden in p1 produced 25 heads this year, but its 99 north counterpart didn’t have a single flowering plant.
Additionally, we did visit p5 this year and found several flowering (and some basal) plants, but were instructed not to harvest them, so we didn’t.
cg heads in the seed dryer prior to going through the ace process in the lab. It isn’t every year when almost all of your gBags fit in the seed dryer at once! I bet I could’ve made em all fit if I really squeezed them, but what’s the rush?
We still have a couple of gBags from 2023 to clean, plus 10 from this year. We like keeping our volunteers stocked! We’ll send exPt01 2024 through first, just after the remnant heads. And so the cycle continues!
Every year since 1996, Team Echinacea members record flowering phenology, taking measuring data and harvest heads of thousands of Echinacea angustifolia plants in common garden experiments. These experimental plots are prairie restorations and abandoned agriculture fields that are managed as grassland habitat. Some plots have multiple ongoing experiments within. Currently, the Echinacea Project currently has 10 established experimental plots.
This project status report will contain updates on experimental plots experimental plots 1, 2, 4, 5 and 8. Reports for the remaining experimental plots can be found on separate posts including Amy Dykstra’s plot (exPt03), the hybrid plots (exPt06, exPt07, exPt09), and the West Central Area common garden (exPt10).
Jo, Emma, Kennedy and Sophia measuring in the picturesque views of exPt02.
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). There are also a number of smaller experiments in it, including fitness of Hesperostipa spartea, aphid addition and exclusion, and pollen addition and exclusion (the latter two experiments were continued the summer of 2022 and will have separate update posts). In 2022, we visited 7,273 of the original 10,673 positions planted and found 2,985 alive. Only 1,111 plants were classified as “flowering” in exPt01 this year. This is a drastic increase from the mere 79 plants that flowered in summer 2021– coincidentally, the plot was burned in the spring prior to summer 2022 and not prior to summer 2021. In summer 2022, we harvested 1,588 total Echinacea heads in exPt01. No additional staples were added to the experimental plot this year.
Some numbers for experiments within exPt01
Inb1: 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. In 2022, of the original 557 plants in INB1, 92were still alive. Of the plants that were alive this year, 40 of them were flowering; this is a drastic difference from summer of 2021 where only one of the plants was flowering.
qgen: The qGen1 (quantitative genetics) 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”). 1,467 plants in qGen1 were alive in 2022. Of those plants, 592 flowered this summer.
Other plots:
exPt02: 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, or 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 2022, we visited 1,856 positions of the 3,961 positions originally planted. We measured 1,438 living plants, of which 651 were flowering, with a total of 1,168 flowering heads. In the fall, we harvested 558 heads from exPt02. We began harvesting on August 10th and completed harvesting on September 12th. The large difference between the number of heads and the number harvested has to do with high levels of seed predation, mainly by ground squirrels. This year, Will, Jennifer, and other members of Team Echinacea published a paper in the American Journal of Botany using data from exPt02 – check it out here! Location: Hegg Lake WMA
exPt04: Experimental plot 4 was planted to gauge whether Echinacea from small remnant populations could be genetically rescued via an outcross to larger, more genetically diverse populations. Caroline Ridley and other members planted this plot in 2008. We did not visit exPt04 this year. Location: Hegg Lake WMA
exPt05: The only experimental plot planted at Staffanson Prairie Preserve (SPP), exPt05, was planted to compare progeny of maternal plants from burned and unburned sections of SPP. There were 2800 plants planted originally, but high mortality made it impractical to visit the plot row-by-row. Now, we and 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 2022 we found 11 living plants, four of which were flowering! There were two heads that should’ve been harvested, but Alex and I forgot to harvest them (oops). Location: Staffanson Prairie Preserve
exPt08: 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 maternal parents of 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. A total of 3,253 seedlings were originally found, but only 363 plants were found alive in 2022. There were 14 flowering plants in 2022, and 15 heads. On a side note, 3 additional flowering plants were found in t-plot, but the heads were eaten before we could harvest any of them. Location: Wagenius property
Start year: Differs between experiment, see above. First ever experimental plot was in 1996.
Location: Differs between experiment, see above.
Overlaps with: …everything!
Data collected: Raw measuring data can be found in cgData repository. Processed data will be eventually uploaded to SQL database. Currently, SQL database has measuring data up until 2021.
Samples or specimens collected: See above for total harvested heads in each plot.
Products: Many publications and independent projects.
Since 1996, members of Team Echinacea have walked, crawled, and ~sometimes~ run next to rows of Echinacea angustifolia planted in common garden experiments. Although protocol varies depending on the experimental plot, every year team members record flowering phenology data, measuring data, and harvest the heads of the thousands of plants we have in common garden experiments.
Note that these experiments are not really gardens. “Common garden” refers to the experimental design. Most of our experimental plots are prairie restorations, a few are abandoned ag fields that are manged as grassland habitat. Some plots have multiple experiments within. The Echinacea Project currently has 10 established experimental plots:
exPts01-10. To avoid repetitiveness of reports on yearly phenology, measuring, and harvesting, this project status report will include updates on all experimental plots 1, 4, 5, and 8. Reports for the others will be elsewhere: Jennifer’s plot (exPt02) Amy Dykstra’s plot (exPt03), the hybrid plots (exPt06, exPt07, exPt09), and the West Central Area common garden (exPt10).
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). There are also a number of smaller experiments in it, including fitness of Hesperostipa spartea, aphid addition and exclusion, and pollen addition and exclusion. In 2021, we visited 6,673 of the original 10,673 planted and found 3,085 alive. Only 79 plants were classified as “flowering” in exPt01 this year. This is a drastic decrease from the 484 plants that flowered in summer 2020– perhaps it is a testament to the benefits of controlled burning (we burned in spring 2020 but not in 2021). In summer 2021, we harvested 77 total Echinacea heads in exPt01 (we have not finished the inventory process). In the fall, we added 403 staples to positions where plants were gone for three straight years. We also converted the flaple>s (pin flags bent in half in spots where we ran out of staples) into regular old staples. There are no more flaples in p1!
exPt04: Experimental plot 4 was planted to gauge whether Echinacea from small remnant populations could be genetically rescued via an outcross to larger, more genetically diverse populations. Caroline Ridley and other members planted this plot in 2008. We did not measure exPt04 this year, but there were no flowering plants. Location: Hegg Lake WMA
exPt05: The only experimental plot planted at Staffanson Prairie Preserve (SPP), exPt05, was planted to compare progeny of maternal plants from burned and unburned sections of SPP. There were 2800 plants planted originally, but high mortality made it impractical to visit the plot row-by-row. Now, we and 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 2021 we found 11 living plants, three of which were flowering! There was only twohead to collect, one head was toppled. Location: Staffanson Prairie Preserve
exPt08: 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 maternal parents of 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. A total of 3,253 seedlings were originally found, but only 443 plants were found alive in 2021. There were 28 flowering plants in 2021, and 32 heads. This is the most flowering heads in p8 we have ever seen, the first year plants flowered in p8 was 2019 and in 2020 five plants flowered. Note that there were an additional 14 heads collected from transplant plot.
Plot management: To ensure that the common garden environment is as similar as possible to the prairie environment we must actively manage it. This management includes removing foreign species and supplementing with natives. One of our main management methods is through fire. We were able to burn burn p8 this spring and hope to burn p1 this spring. We also collected seed to spread after burns including Schizachyrium scoparium, Bouteloua curtipendula, along with multiple species of Solidago and quite a few Asters.
Asclepias viridiflora in p1: In 2019, plugs of an uncommon prairie milkweed, Asclepias viridiflora, were planted in Experimental plot 1. The purpose of this experiment is to assess the survival and fitness of A. viridiflora. Assessing vitality will also provide a frame of reference for species conservation across modern prairies. We did not assess the A. viridiflora plants this year.
Hesperostipa demography: In 2009 and 2010, porcupine grass (Hesperostipa spartea, a.k.a. “stipa”) was planted in experimental plot 1. In total, 4417 seeds were planted, 1 m apart from each other and all 10 cm north of Echinacea plants. Between 2010 and 2013, each position was checked, and the plant status recorded. Since 2014, we have searched for flowering plants. The data from this summer can be found here cgdata/summer2021/stipaSearch & Dropbox/CGData/Stipa/225_measure/measure2021 in addition to data in these locations there may be paper data sheets that may or may not have been entered yet. These data have not been processed yet.
Inb1: 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. We continued to measure fitness and flowering phenology in these plants. In 2021, of the original 557 plants in INB1, 95were still alive. Of the plants that were alive this year, there was only one flowering plant. All individuals were planted in 2001.
qgen: The qGen1 (quantitative genetics) 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”). In 2021 we assessed survival and fitness measures of the qGen1 plants. 1,519 plants in qGen1 were alive in 2021. Of those, 2.4% flowered in 2021, this is down from 17% last year. All were planted in 2003.
Team moral: Measuring experimental plot 1 is a large task for the team, it takes many days, usually hot days, with a project as large as this it is hard for the team to feel a sense of progress each day. So this year in hopes to keep the team more motivated I(Mia) took two strategies. First, we constructed a physical progress tracker, each square represented a segment in the plot A segment is a section of a row, rows are too big so we break them into sections for ease of measuring. After each day of measuring the team was able to count how many segments they completed and color in each square. The team had fun with their square decoration and got creative with it, there was various themes, fruits, names for peoples last days etc. I also made maps of the plot for each day of measuring that were color coded with what positions have been measured and which ones haven’t. Lastly, I do believe I may have bribed the team with cookies. Having multiple different forms of motivation, really kept the team motived and made the project less daunting. It was also added some excitement into measuring p1, a task that isn’t known for being too exciting.
one of the progress maps purple is done green is not done
one of the progress maps purple is done green is not done
A hearty crew finishing measuring p1 during one of the only rainy days of the season
Progress tracker aka Gopher tunnel of love
For more information on survival in common garden experiments, see this flog post about survival in common gardens.
Start year: Various, see individual listings above. First ever planting was 1996.
Location: Various, see above
Overlaps with: Pretty much everything we do.
Data/ materials collected: Measure data for all plots. All raw measure data available in cgData repository. Processed data should eventually be available in SQL database; ask GK for status of SQL database.
new p79 stake file: Jared made a cleaned up stake file for p7 and p9 that can be found here
p2 stake file/points shot: Amy shot points in p2 this year the points can be found in these 3 jobs
Last week Stuart, Gretel, Jared, and I headed northward from Chicago to Minnesota to perform the first prescribed burn of the season! On our drive up we hit some snow that was almost whiteout conditions very exciting, especially for April. We arrived in Douglas County late Wednesday night and quickly bundled into our sleeping bags.
Stuart examining the test fire of the Island
The next morning, we walked the unit/p8 and saw two bald eagles flying over the plot, we decided that this was a good omen for the burn. There are two areas in the unit that haven’t been burned in the past that we decided to burn this year, this was the island that is northeast of the plot and the “bee trees”. After examining the unit we set off to prep the unit and gather supplies. After a break for lunch, we ran a test of how the wooded area would burn by burning the island area. This burn went well, the fire moved slowly but we did kill a frog :(. Halfway through this burn Ruth and Frank arrived from the Cities, they were greeted with excitement and backpack sprayers.
After the success of burning through the woods in the island, we decided to burn through the bee trees. The bee trees burned very slowly Frank and I spend most of the burn focused on ensuring that no sparks from the bee’s trees got taken in the wind downhill. To the south of the bee trees, the burn brake is only mowed and still has quite a bit of brome that could be fuel. We were all shocked by the civilized behavior that the fire had around this burn break. Once we had a sufficient backfire Stuart light the head fire in the windward portion of the plot and boy it was spectacular. Our civilized fire politely ripped through the brome of p8 and even left many pin flags untouched!
The backfire heading southThe head fire heading through p8
After we were satisfied with the large p8 unit fire being out we gathered, including John VanKempen who arrived during the course of the p8 burn. We then headed down to Jean’s prairie plant garden and Jared, who was the burn boss for the final two burns, light a nice line around the perimeter of the garden, this burn only took 16 minutes. When we were waiting for the garden to burn we noticed a small adjacent patch of dried duff and we decided to burn that too! This burn went even faster than the prairie garden it was also much more powerful.
The burn crew! Left Gretel, Jared, Mia, John, Ruth, Frank (Stuart not pictured)
After the prairie garden and adjacent area were done burning, Stuart, Gretel, Jared and I made sure that everything was put out back at p8. We found a smoldering log and made some s’mores!
The next day, Jared and I inspected the remnant sites that we are planning on burning. Jared, Stuart, and I also broadcast some native prairie seeds (mostly two grasses: side-oats grama and little bluestem) that we collected in the fall in p8.
We performed this burn a week ago now, on Earth Day, performing a prescribed burn that aids in the restoration of the prairie was a wonderful way to celebrate. This was my first prescribed burn, overall it was really fun, impressive, exciting, and also boring. I am very excited to be heading back up to Minnesota to conduct more burns but also to see how the community changes after the burn.
The charred and snow-sprinkled plot on Saturday morning
Since 1996, members of Team Echinacea have walked, crawled, and ~sometimes~ run next to rows of Echinacea angustifolia planted in common garden experiments. Although protocol varies depending on the experimental plot, every year team members record flowering phenology data, measuring data, and harvest the heads of the thousands of plants we have in common garden experiments.
Note that these experiments are not really gardens. “Common garden” refers to the experimental design. Most of our experimental plots are prairie restorations, a few are abandoned ag fields that are manged as grassland habitat. Some plots have multiple experiments within. The Echinacea Project currently has 10 established experimental plots:
exPts01-10. To avoid repetitiveness of reports on yearly phenology, measuring, and harvesting, this project status report will include updates on all experimental plots 1, 2, 4, 5, and 8. Reports for the others will be elsewhere: Amy Dykstra’s plot (exPt03), the hybrid plots (exPt06, exPt07, exPt09), and the West Central Area common garden (exPt10).
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). There are also a number of smaller experiments in it, including fitness of Hesperostipa spartea, aphid addition and exclusion, and pollen addition and exclusion. In 2020, we visited 4,340 of the original 10,622 planted and found 3,258 alive. Only 484 plants were classified as “flowering” in exPt01 this year. This is a drastic increase from the nearly 70 plants that flowered in summer 2019 – perhaps it is a testament to the benefits of controlled burning (we burned in spring 2020 but not in 2019). In summer 2020, we harvested ~815 total Echinacea heads in exPt01 (I have not finished the reconciliation process). In the fall, we added 484 staples to positions where plants were gone for three straight years, however, we ran out of staples, so 130 of these positions have “flaples” which are bent pin flags.
exPt02: Heritability of flowering time is the name of the game in exPt02. Planted in 2006, exPt02 was planted to assess heritability of flowering start date and duration in Echinacea. In summer 2020, we visited 2,010 positions of the 3,961 positions originally planted. We measured 1,638 living plants, of which 444 were flowering. In the fall, we harvested ~626 heads from exPt02. We do not have an exact number of heads harvested from exPt02 yet, as we have not had time to complete head reconciliation. Location: Hegg Lake WMA
exPt04: Experimental plot 4 was planted to gauge whether Echinacea from small remnant populations could be genetically rescued via an outcross to larger, more genetically diverse populations. Caroline Ridley and other members planted this plot in 2008. We did not measure exPt04 this year. Location: Hegg Lake WMA
exPt05: The only experimental plot planted at Staffanson Prairie Preserve (SPP), exPt05, was planted to compare progeny of maternal plants from burned and unburned sections of SPP. There were 2800 plants planted originally, but high mortality made it impractical to visit the plot row-by-row. Now, we and 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 2020 we found 11 living plants, two of which were flowering! There was only one head to collect, since one of the flowering plants exhibited only vertical development (no head). Location: Staffanson Prairie Preserve
exPt08: 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 maternal parents of 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. A total of 3,253 seedlings were originally found, but only 562 plants were found alive in 2020. There were 5 flowering plants in 2020, and 5 heads. Note that there were an additional 2 heads collected from transplant plot.
Plot management: To ensure that the common garden environment is as similar as possible to the prairie environment we must actively manage it. This management includes removing foreign species and supplementing with natives. One of our main management methods is through fire. We were unable to burn plots this fall however we hope to burn p8 and p1 this spring. We also collected seed to spread after burns including Schizachyrium scoparium, Bouteloua curtipendula, along with multiple species of Solidago and quite a few Asters.
Asclepias viridiflora in p1: In 2019, plugs of an uncommon prairie milkweed, Asclepias viridiflora, were planted in Experimental plot 1. The purpose of this experiment is to assess the survival and fitness of A. viridiflora. Assessing vitality will also provide a frame of reference for species conservation across modern prairies. In 2020 a protocol was developed to identify and measure A. viridiflora. These data are waiting to be entered and analyzed.
Hesperostipa demography: In 2009 and 2010, porcupine grass (Hesperostipa spartea, a.k.a. “stipa”) was planted in experimental plot 1. In total, 4417 seeds were planted, 1 m apart from each other and all 10 cm north of Echinacea plants. Between 2010 and 2013, each position was checked, and the plant status recorded. Since 2014, we have searched for flowering plants. The data from this summer can be found here cgdata/summer2020/stipaSearch, these data have not been processed yet.
Inb1: 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. We continued to measure fitness and flowering phenology in these plants. In 2020, of the original 557 plants in INB1, 111 were still alive. Of the plants that were alive this year, 30 (27%) were flowering. This is up from the 4% that were flowering last year. All individuals were planted in 2001.
qgen: The qGen1 (quantitative genetics) 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”). In 2020 we assessed survival and fitness measures of the qGen1 plants. 1,642 plants in qGen1 were alive in 2020. Of those, 17% flowered in 2020. All were planted in 2003.
Stuart and John measure Echinacea plants in exPt02 under a gloomy sky
For more information on survival in common garden experiments, see this flog post about survival in common gardens.
Start year: Various, see individual listings above. First ever planting was 1996.
Location: Various, see above
Overlaps with: Pretty much everything we do.
Data/ materials collected: Measure data for all plots. All raw measure data available in cgData repository. Processed data should eventually be available in SQL database; ask GK for status of SQL database. GPS points were shot for the exPt09 flowering plant. Find the GPS jobs containing the exPt09 corners here: Dropbox/geospatialDataBackup2020/convertedASVandCSV2020/EXPT9_20200917_DARW.asv The stake file to find exPt5 plants is here: Dropbox/geospatialDataBackup2020/stakeFiles2020/exPt05stakeFile.csv Products: Many publications and independent projects.
Team Echinacea established quantitative genetics experiments to determine the additive genetic variance of fitness in Echinacea, with the idea that we can estimate evolutionary potential of study populations. Quantitative genetics experiments 2 and 3 (qGen2 and qGen3) represent the third generation of Echinacea in our common garden experiments. The grandparents of qGen2 and qGen3 are the 1996 and 1997 gardens. Plants from these experiments were crossed to generate qGen1 (a.k.a. Big Batch), and plants in qGen1 were crossed to produce seed for qGen2 and qGen3, which now inhabit exPt8.
We visit exPt8 every year to assess fitness of Echinacea in the plot. Originally, 12,813 seeds were sown in the common garden. Seeds from the same maternal and paternal plant were sown in meter-long segments between nails. A total of 3253 seedlings were originally found, but only 669 plants were found alive in 2019.
Jay, John, and Avery assess fitness of young Echinacea in exPt8. They’re so tiny (the Echinacea, that is… Jay, John, and Avery are regular sized).
In an exciting turn of events, we found a flowering plant in qGen2 this year! This was the first flowering plant found in exPt8. Fortunately for our one flowering plant, it had four flowering friends to cross with from the Transplant Plot. We took phenology data on the qGen2 head, measured it, and harvested it.
The presence of a flowering plant influenced Riley Thoen to make a new measuring form for exPt8 in 2020. In the past, the exPt8 measuring form was very different from other measuring forms. Through 2019, we measured all leaves of basal plants in exPt8; we only measure the longest basal leaf in other plots. Riley designed the 2020 exPt8 measuring form to mirror the measuring forms from other common gardens. In the future, the exPt8 measure form will have a head subform and team members will only have to measure the longest basal leaf of each plant found.
Start Year: 1996 and 1997 (Grand-dams), 2003 (qGen1 – dams), 2013 and 2015 (qGen2 and qGen3, respectively)
Data/material collected: phenology data on the flowering plant and transplant plot plants (available in the exPt1 phenology data frames in the cgData repo), measure data (cgData repo), and harvested heads (data available in hh.2019 in the echinaceaLab package; heads in ACE protocol at CBG).
Last Friday, I was dispatched by Stuart to find the number of plants/ achenes planted in each experimental plot, along with the number alive as of a recent year (2017-2019, based on the plot). Although records of some plots were a bit harder to come across that others, I was able to compile data from each plot (besides p10 – planted 2019 – data coming soon). This would not have been possible without the help of Gretel, so thanks GK! I have attached a small datasheet with the survival data.
In the history of the Echinacea Project, the team has sown 31,888 Echinacea viable achenes in experimental plots. There were many more sown that likely did not have a seed. Team members found 3634 seedlings from these seeds, not including Amy D’s experimental plot 3 and remnant seedling refinds. The team has planted 18,869 Echinacea seedlings in experimental plots, not including p10 – planted at West Central Area HS in 2019. Finally, 7090 Echinacea are currently alive in the experimental plots!
Alex and Tracie search for juvenile Echinacea plants in experiments qGen2 & qGen3.
In 2017, we found 1006 three-year-old plants out of the 2526 original seedlings found in 2014 (we found 1724 plants in 2016) in the qGen2 cohort. In the qGen3 cohort we found 248 of the 644 seedlings.
The main goal of the qGen2 and qGen3 experiments is to quantify the evolutionary potential of two remnant prairie populations of Echinacea angustifolia by estimating the additive genetic variance of fitness. We make estimates for two mating scenarios. The first scenario is an experimental crossing design with all matings among plants from two “core” sites: SPP and LF (core x core). The second design uses sires (pollen donors) from the core and dams from sites peripheral to the core. The crosses performed (core x core, core x periphery) in this experiment will quantify additive genetic variance for fitness in each site and each experimental group. Additionally, we will test for differentiation among families; do progeny from sires differ after accounting for maternal (dam) effects?
Comparing germination and first year survival between the qGen2 & qGen3 cohorts:
exp
approxFullAcheneCt
totalAcheneCt
seedlingCt
germination
firstYrSurvival
qGen2
6300
26144
2581
41%
84%
qGen3
6200
19777
644
10%
38%
Start year qGen3: 2015
Start year qGen2: 2013
Location: The sires (pollen donors) are in the remnants Landfill and Staffanson. The dams (seed plants) are in exPt 1 and they originate from remnants. Specifically, the grand-dams (seed plants of dams) are from remnants Landfill (core) & around Landfill (peripheral) and remnants Staffanson (core) & railroad crossing sites (peripheral). All progeny are in exPt 8.
Overlaps with: Heritability of fitness–qGen1
Data collected: We used handheld computers to collect data on juvenile plants.
You can find more information about Heritability of fitness–qGen2 & qGen3 and links to previous flog posts regarding this experiment at the background page for the experiment.
Alex and Lea measure qGen3 seedlings with much enthusiasm.
The main goal of the qGen2 and qGen3 experiments is to quantify the evolutionary potential of two remnant prairie populations of Echinacea angustifolia by estimating the additive genetic variance of fitness. We make estimates for two mating scenarios. The first scenario is an experimental crossing design with all matings among plants from two “core” sites: SPP and LF (core x core). The second design uses sires (pollen donors) from the core and dams from sites peripheral to the core. The crosses performed (core x core, core x periphery) in this experiment will quantify additive genetic variance for fitness in each site and each experimental group. Additionally, we will test for differentiation among families; do progeny from sires differ after accounting for maternal (dam) effects?
In 2016, we found 1724 two year old plants out of the 2581 locations where plants had previously been found for the qGen2 cohort and 644 seedlings in the qGen3 cohort.
Comparing germination between the qGen2 & qGen3 cohorts:
exp
approxFullAcheneCt
totalAcheneCt
seedlingCt
germination
qGen2
6300
26144
2581
41%
qGen3
6200
19777
644
10%
Our crossing success, measured by the proportion of full achenes to total achenes crossed, increased in qGen3 (31%) compared to qGen2 (24%). While we planted approximately the same number of full achenes in the qGen2 & qGen3 cohorts, the germination rate was 4 times greater in qGen2 (41%) compared to qGen3 (10%). This difference was likely due to differences in environmental conditions. The Spring of 2016, was quite dry and probably tough on Echinacea seeds and sprouts.
Start year qGen3: 2015
Start year qGen2: 2013
Location: exPt 1 (dams), remnants Landfill and Staffanson (sires), remnants Landfill (core) & around Landfill (peripheral) and remnants Staffanson (core) & railroad crossing sites (peripheral) (grand-dams), exPt 8 (progeny)
Overlaps with: Heritability of fitness–qGen1
Data collected: We used handheld computers to collect data on seedlings and juvenile plants.
You can find more information about Heritability of fitness–qGen2 & qGen3 and links to previous flog posts regarding this experiment at the background page for the experiment.