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 1, 2, 4, 5 and 8, as well as management updates for all plots. Specific 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).
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 last experiment was continued the summer of 2023 and will have separate update posts). In 2023, we visited 3,699 of the 10,992 positions planted and found 3,118 alive. 560 plants were classified as “flowering” in exPt01 this year. This is a little less than half of the plants that flowered in summer 2022 (1,111) – an interesting note considering exPt01 was burned the spring of both 2022 and 2023. In summer 2023, we harvested 796 total Echinacea heads in exPt01. We also added 270 additional staples to the experimental plot this year, signifying positions were a living plant has not been found for over three years.
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 2023, of the original 557 plants in INB1, 85 were still alive. Of the plants that were alive this year, 24 of them were flowering; this count is down from summer of 2022 where 40 of the plants were 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”). 1,417 plants in qGen1 were alive in 2023. Of those plants, 298 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 2023, we visited 1,855 positions of the 3,961 positions originally planted. We measured 1,283 living plants, of which 118 were flowering, with a total of 148 flowering heads. In the fall, we harvested 67 heads from exPt02. 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. Last 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 2023 we found 11 living plants, none of which were flowering! We did locate one new flowering plant within the plot boundaries. 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 385 plants were found alive in 2023. There were 15 flowering plants in 2023, and 15 heads. On a side note, one additional flowering plant was found in t-plot, and we harvested three heads from that. Location: Wagenius property
Experimental plot management:
Burned exPt01 (3 May 2023) and exPt08 (17 May 2023)
Replanted pedicularis in exPt01 and exPt10, augmenting Drake’s experimental treatments (replaced ones that died)
Broadcast seed in p8 after the spring burn and in the fall, including Comandra umbellata
Stuart trimmed flowering A. gerardii and S. nutans just north of tplot
Some plots in hegg (not exPt02) got run over by heavy machinery
We did not:
Treat sumac
Weed in exPt01 (except hawkweed)
Treat ash in exPt08, but we noticed that ash south of plot responded favorably to last year’s treatment
Details
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 uploaded to SQL database. Currently, SQL database has measuring data up until 2022.
Samples or specimens collected: See above for total harvested heads in each plot.
Products: Many publications and independent projects.
Harvesting Echinacea heads in the common garden experiments this fall was quite the adventure! During the last week in June, the crew got a taste of harvesting when decapitating non-native Echinacea pallida at Hegg before they could produce seeds. Lobbing off Echinacea heads with wild abandon was quite the thrill after spending a month conscientiously navigating the common gardens and measuring plants with care. In total, we cut approximately 824 heads off of 224 flowering pallida plants.
To harvest or not to harvest? Here is Daytona before removing a pallida head.
The field crew started harvesting heads to be cleaned in exPt02 on August 10th. Our final day of harvesting in exPt02 was September 12th, and during that month we harvested exactly 480 Echinacea heads. Our bountiful harvest in exPt02 was thwarted by the local population of thirteen-lined ground squirrels. Many of this summer’s field team recounts witnessing these rodents ruthlessly rip the head off an Echinacea, look them straight in the eye, and run away. In the end, our 480 heads accounted for only 41% of the heads we planned on harvesting prior to the squirrel shenanigans.
This was a typical view while harvesting in exPt01! So much tall Sorghastrum and Andropogon at eye level.
We also harvested heads in some sites with fewer flowering Echinacea. In exPt08, we harvested 8 heads, and there were three flowering plants in t-plot that may have succumbed to the ground squirrels before we could harvest them. In exPt07 and exPt09, we harvested a combined total of 130 heads. The mysterious exPt05, which required a GPS to be found at Staffanson Prairie, had 4 flowering plants with 2 heads that should’ve been harvested, but didn’t (oops!).
The harvest of our largest experimental plot, exPt01, began on Aug 30th. The exPt01 madness did not end until our final four heads were harvested on October 10th by Gretel and Stuart, which is the latest harvest recorded in Echinacea Project history! The grand total for number of heads harvested in exPt01 was a whopping 1,494 heads.
Manogya hard at work in exPt01 (photo credits to Alex).Teamwork makes the dream work (another great photo from Alex)!
Between all of our common garden experiments, we harvested a total of 2,112 heads!
The inbreeding 2 experiment was planted in exPt1 in 2006 to determine how genetic drift is differentially affecting average fitness of remnant populations. In 2005, team members crossed common garden plants from seven remnant populations. There are three cross types: inbred (crossed to a half-sib; I), within population (randomly chosen; W), and between population (B). Each year, team members assess flowering phenology and fitness of Echinacea in the inb2 common garden.
In 2019, the team searched for Echinacea at 508 positions of the original 1443 positions planted in inb2. In total, we found 351 living plants. Four plants flowered in 2019 but only three produced achenes. Since 2006, 163 Echinacea in inb2 have flowered; they have produced a total of 336 flowering heads.
This winter, Riley Thoen is working on analyzing data and drafting a manuscript for inb2. In these endeavors, he found a small discrepancy in inb2 data: not all plants that were planted in the inb2 plot have a complete pedigree. Therefore, only a subset of the total can be used for analysis. A total of 1136 plants with a complete pedigree were planted in inb2, and of those, 277 were found alive in 2019. All four plants that flowered in 2019 have known pedigrees. A total of 138 plants of known pedigree have flowered and they have produced 284 total heads since the plot was planted in 2006. Surprisingly, within-remnant crosses have the lowest survival of all cross types, at 20%. Inbred crosses have 24% survival and between-remnant crosses have 30% survival. Riley is starting to push data analysis forwards and will certainly post updates on the flog when more discoveries are made!
Data/material collected: flowering phenology on the flowering 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).
Products:
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
Reina, Pamela, and Mike with the photosynthesis machine used in Kittelson et al. (2015)
In 2017, of the original 1,470 plants in INB2, 473 were still alive. Of the plants that were alive this year, only 4 were flowering (0.8%) and 71% have never flowered. Interestingly, there are differences in survival for each cross level. 31% of the most inbred plants were living in 2017, compared to 39% of the plants in the between-remnant cross level. All individuals were planted in 2006.
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. We continued to measure fitness and flowering phenology in these plants.
Overlaps with: Phenology and fitness in P1, Inbreeding experiment–INB1
Physical specimens: We harvested 2 heads from INB2 that will be processed in the lab with other heads harvested from P1.
Data collected: We used handheld computers to collect fitness data on all plants in INB2.
Products: The below papers were published in summer 2015:
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
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
You can find more information about the Inbreeding experiment–INB2 and links to previous flog posts regarding this experiment at the background page for the experiment.
Reina, Pamela, and Mike with the photosynthesis machine used in Kittelson et al. (2015)
In 2016, we continued the INB2 experiment to investigate 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. We continued to measure fitness and flowering phenology in these plants.
This year, of the original 1,470 plants in INB2, 557 were still alive. Of the plants that were alive this year, 2% were flowering and 75% have never flowered.
Overlaps with: Phenology and fitness in P1, Inbreeding experiment–INB1
Physical specimens: We harvested 9 heads from INB2 that will be processed in the lab with other heads harvested from P1.
Data collected: We used handheld computers to collect fitness data on all plants in INB2.
Products: The below papers were published in summer 2015:
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
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
You can find more information about the Inbreeding experiment–INB2 and links to previous flog posts regarding this experiment at the background page for the experiment.
Reina, Pamela, and Mike with the photosynthesis machine used in Kittelson et al. (2015)
In 2015, we continued to study the effects of inbreeding on Echinacea angustifolia fitness. This experiment was planted in 2006 where each plant was produced from one of three cross types: between maternal half siblings; between plants originating from the same remnant, but not sharing a maternal parent; and between individuals from different remnants. We continued to measure fitness and flowering phenology in these plants.
This year, of the original 1443 plants in INB2, 561 were still alive. Of the plants that were alive this year, 8.3% were flowering and 76.3% have never flowered – we’re still waiting! Among the plants that were flowering, mean head count was 1.53 heads, with a maximum of five heads.
Products: Fitness measurements were collected during our annual assessment of fitness in P1.
The following paper was published in summer 2015 based on fieldwork conducted in 2013.
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
Experimental plot 1 (P1) encompasses 11 different experiments originally planted with a total of 10673 Echinacea individuals. These experiments include long-term studies designed to compare the fitness of Echinacea from different remnant populations (“EA from remnants in P1”), examine the effects of inbreeding on plant fitness (“INB” and “INB2”), and explore other genetic properties of Echinacea such as trait heritability (“qGen”). In 2014, Team Echinacea measured plant traits for the 5409 Echinacea plants that remain alive and followed the daily phenology of 567 flowering heads. Echinacea began producing florets on July 1 and continued flowering in P1 until August 24. The data collected in 2014 will allow us to estimate the heritability of various traits and assess the lifetime fitness of plants from the numerous experiments.
Today was a very nice day today. There were clear skies, it was sunny, there was just enough wind to keep the mosquitos at bay, but no pollinators. Up at Hegg Lake the wild Monarda have started blooming in force, along with the swamp milk weed (which is shown below).
In the morning Sara and Sarah went out to work on their individual projects, while Lydia, Marie, Gretel, Ilse, Per and Hattie worked on phenology.
In the afternoon Gretel, Lydia and Ilse worked on the QGen- which meant continuing the pollenating that has been going on since flowering began. Myself, Sarah, Sara, and Marie started and finished measuring the echinacea in the INB2 garden! Woot!
It was another hot day on the prairie front. We all broke off into little groups today to take care of business. This morning Reina, Mike, and Pam assed herbivory in INB1 and INB2. Kory, Marie, Sarah B, and Dayvis were all scattered about working on their independent projects. Lydia and Sara Z assed the maternal plants in the common garden that are to be involved in the crosses of the quantitative genetic experiment, and on the other side of things Gretel, Shelley (Gretel’s Mother), Ruth and I went to Landfill Core to gather pollen from the sires that are to be involved in the crossing experiment. We used toothpicks to scrape off the pollen into vials that will be used for pollenating the maternal plants in the common garden. We were able to get pollen from about 40 of 55 plants at the site. The 15 others were just a little too immature to collect pollen from today.
This afternoon encompassed an hour of stipa searching for some and then continuation of some independent work on projects. Herbivory assessment continued as well. Stuart, Gretel, Ruth and I went for a nice long walk around SPP. We walked a U-shaped transect and used a randomized scheme for assigning sires to be used in the quantitative genetics experiment. We caged/bagged heads that were to be used and gps-ed the plants.
Having dodged the misfortune of convening on Friday the thirteenth, Team Echinacea had a fruitful day in the field.
The morning was occupied by independent projects. Kory, Jennifer, and Jon went to CG2 at Hegg Lake to look for pollinators visiting flowering Echinacea. Dayvis and Marie also went to Hegg Lake to gather data for their respective projects. Sarah B. once again visited multiple remnants to monitor flowering dates. Meanwhile, Lydia, Ilse, and Gretel assessed phenology and flowering head count in the Common Garden and ’99 South Garden to gear up for this summer’s pollination experiment.
In the afternoon, Pam and Reina finished taking measurements of plants in INB2 for their study. The progress of their work benefited greatly from the recent acquisition of a Red Flyer wagon. Lydia (shown below) braved the wrath of protective ants in her quest to acquire blue aphids for her addition/ exclusion experiment. The rest of the team returned to CG2 to finish measuring plant fitness traits.
After a long day’s work, Dayvis cooked up a delicious meal of split pea soup and arepas (Venezuelan corn cakes). Many crew members are experiencing discomfort due to chigger bites, although some are more prone to vocalize their displeasure than others.