2020 Update: Flowering phenology in the remnants

In 2020, we collected data on the timing of flowering for 855 flowering plants (1071 flowering heads) in 31 remnant populations. The plants ranged from having 1 to 8 flowering heads. The earliest bloomers initiated flowering on June 22nd . Plant 22195 at NWLF was the latest bloomer, only beginning to shed pollen on September 14th, nearly a month after the second-latest flowering plant had ceased producing pollen (August 18th). As is typical for the latest bloomer of a season, township mowers had mowed over this plant earlier in the season, which is perhaps why it took longer for it to sprout a new flowering stem. Peak flowering was on July 9th, when 886 heads were flowering.

A major part of the motivation behind this year’s effort in monitoring phenology was to collect baseline data on flowering rates and timing. Team Echinacea recently received funding to perform prescribed burns in these populations. Next summer, we will compare flowering patterns in populations before and after fires to understand how burns drive the effects of timing of flowering on mating patterns and fitness of individuals in natural populations.

Start year: 1996

Location: Roadsides, railroad rights of way, and nature preserves in and around Solem Township, MN

Overlaps with: phenology in experimental plots, demography in the remnants, gene flow in remnants, reproductive fitness in remnants

Data/materials collected: We identify each plant with a numbered tag affixed to the base and give each head a colored twist tie, so that each head has a unique tag/twist-tie combination, or “head ID”, under which we store all phenology data.We monitor the flowering status of all flowering plants in the remnants, visiting at least once every three days (usually every two days) until all heads were done flowering to obtain start and end dates of flowering. We managed the data in the R project ‘aiisummer2020′ and will add the records to the database of previous years’ remnant phenology records, which is located here: The dataset is ready to be updated, but I don’t believe it has been at the time of writing.

A flowering schedule for individuals from all remnants. Notice the gap between when second-to-last flower ceased pollen production and when the latest bloomer began on September 14th!

A flowering curve (created here using the R package mateable) summarizes the flowering phenology data that we collected in 2020, indicating the number of individuals flowering on a given day and the flowering period for all individuals over the course of the season.

We shot GPS points at all of the plants we monitored. Soon, we will align the locations of plants this year with previously recorded locations and given a unique identifier (‘AKA’). We will link this year’s phenology and survey records via the headID to AKA table.

You can find more information about phenology in the remnants and links to previous flog posts regarding this experiment at the background page for the experiment.

Products: A dataset of flowering phenology is ready to be posted on the website. It is currently located in Dropbox\remData\105_assessPhenology\phenology2020\phen2020_out and is available upon request. The headIds in this dataset have not yet been merged with the akas (long-term identifiers) in the demography dataset.

2020 Update: ExPt 10 and Gene Flow in the Remnants

During the summer of 2019, Team Echinacea planted over 1400 E. angustifolia seedlings into 12 plots in a prairie restoration at West Central Area High School in Barrett, MN. We planted seedlings from three sources: (1) offspring from exPt1, (2) plants from my gene flow experiment, and (3) offspring from the Big Event. To test how different fire regimes affect fitness in Echinacea, folks from West Central Area plan to apply regular fall burn treatments to four plots, regular spring burn treatments to four other plots, and the remaining four plots will not be burned. I’m not sure if they were able to perform these burns as planned in Fall 2020 given COVID restrictions this spring and fall, but John Van Kempen would be the man to ask about that. I believe they were able to do the burns in the spring.

This summer, the team measured the 1-year old seedlings from my gene flow study in exPt10, as well as a few seedlings from the other plantings within the plot. The seedlings from my gene flow experiment are the offspring of open-pollinated Echinacea in 9 populations in the study area. I am assessing the paternity of these seedlings to understand contemporary pollen movement patterns within and among the remnants. In summer 2018, I mapped and collected leaf tissue from all Echinacea individuals within 800m of the study areas and harvested seedheads from a sample of these individuals (see Reproductive Fitness in Remnants). In spring 2019, I germinated and grew up a sample of the seeds that I harvested to obtain leaf tissue for genotyping.

Then, with the team’s help, I planted these seedlings in exPt10 in June 2019. I also collected seeds and leaf tissue in summer 2019 to repeat this process, but I did not germinate the achenes in the following spring because I was not able to assess seed set due to the broken x-ray machine at the CBG and then COVID-related restrictions. I hope to germinate those this spring and plant in summer 2021. I am working on extracting the DNA from the leaf tissue samples I have, which I will use to match up the genotypes of the offspring (i.e., the seeds) with their most likely father (i.e., the pollen source).

Here are some fun facts about the seedlings we found in exPt 10:

  • The longest leaf we saw was 19 cm! Most were much smaller (see below).
A histogram of seedling leaf lengths, ranging from 1 to 19 cm
  • The leafiest plant we saw had 4 leaves (though one had been munched)
A histogram of leaf counts per plant, which ranged from 1 to 4
  • Overall we found 424 seedlings alive of the 598 that we searched for, or 71%. The ones we didn’t find are probably dead, but we’ll look for them again next year to make sure we didn’t just miss them.

I’m looking forward to seeing these friends again next year.

Allie gives a thumbs after successfully finding a baby Echinacea plant in p10!

Start year: 2018

Location: West Central Area High School’s Environmental Learning Center, Barrett, MN, Remnant prairies in Solem Township, Minnesota

Overlaps with: Reproductive Fitness in RemnantsPhenology in the Remnants

Products: Survival data for seedlings planted in summer 2019 from Amy W’s gene flow experiment, located in the cgdata bitbucket repository.

Team members who worked on this project include: Amy Waananen, John Van Kempen

Demcirc – Entry # 1

Stuart and Drake on November 24th, 2020 discussed how Drake should go about entering the data he collected on his circles, apply named “Drake’s circles for the possible intedended purposes of burning with fire and planting with parasites” or “demcirc” for short.

Drake went to the following sites and set up 10 randomly placed (to the best of his ability using his cellphone, laptop, and paper maps) circles: South of Golf Course, East Riley/Riley (10 total across both), Woody’s, Leuffler’s Corner East, Leuffler’s Corner West, Hill across from Leuffler’s Corner West, RRX, NRRX, In front of the Tower Rd Tower, Nice Island, The hill just east of Unity Drive on Hwy 27, Around Landfill North, Around Landfill South, Sap, TH, NNWLF, and KJs.

Additionally 30 circles each at: Aan, Eelr, Landfill, AgalinisRRX (NW Corner of Roland Lake RD and Hwy 55), and all-around Wennersborg Rd SW & HWY 55.

Each circle had a radius of ~0.4m and I recorded every species present within the circle and I recorded whether they were flowering (or had flowered at some point that year) or not.

Spatial mate-limitation in remnants

During the summer of 2020, I designed an observational study to assess the extent to which reproduction in self-incompatible species is limited by spatial proximity to mates. While many researchers assume mate-limited reproduction is common in prairie species, few studies demonstrate this relationship in species other than Echinacea. I hypothesize that many self-incompatible prairie species produce few viable seeds because of small population size and spatial isolation from mates. To test this hypothesis, I selected 8 common self-incompatible native prairie species and 25 remnant sites within which to work. I quantified isolation in meters from the two nearest flowering conspecifics and collected seeds for more than 1,100 individual plants at 25 prairie remnant sites. I hope to complete lab work over the next 6 months where I will be cleaning and counting viable seeds for each individual plant. Results from this study will provide evidence about how widespread mate-limited reproduction is in self-incompatible prairie species.

Start year: 2020

Location: Douglas County, Minnesota; 25 roadside remnant sites Information about how many species of each species were sampled from each site, check out this file ~Dropbox/teamEchinacea2020/leaRichardson/siteCheckList8.29.20.xlsx

Materials collected: Seed heads collected from over 1,100 plants will be stored at the Chicago Botanic Garden but are currently in my Evanston apartment.

Data collected: Find data related to this project in the aiisummer2020 repository in ~aiisummer2020/plasInRems/leaStuff/

Experimental plot one parasite planting (pipp)

Throughout the summer, I designed and collected materials to establish an experiment in experimental plot 1 to study parasites and their impact on the community of host plants they live in. Parasitic plants are plants which absorb nutrients from neighboring plants. Parasitism is an important part of nutrient cycling in many ecosystems and parasite scientists hypothesize it to be an important part of prairie ecosystem maintenance.

This experiment has three factors, each with two levels (presence or absence), but three factor-level combinations are impossible, because the presence of parasites is confounded with presence of soil. Which translates to me having 216 row x position combinations in which I randomly assigned Comandra umbellata, Pedicularis canadensis, and soil plugs. However, because roots trap soil and therefore soil is always carried in with parasites, the two are confounded and so we used soil transplants to account for this.  

Additionally, in all 216 of my “pipp places” I distributed seeds from 32 native plant species: Achillea millefolium (5 seeds), Allium stellatum (3), Anemone patens (3), Astragalus canadensis (1), Bromus kalmii (5), Calylophus serrulatus (3), Carex brevior (3), Carex gravida (1), Dalea purpurea (5), Delphinium carolinianum (2), Elymus canadensis (1), Elymus trachycaulus (5), Geum trifolium (2), Grindelia squarrosa (2), Heuchera richardsonii (2), Koeleria macrantha (5), Liatris aspera (3), Lilium philadelphicum (3), Mirabalis nyctaginea (1), Monarda fitulosa (3), Muhlenbergia cuspidata (1), Oenothera biennis (3), Oxytropis lambertii (2), Pediomelum argophyllum (1), Scrophularia lanceolata (2), Sisyrinchium campestre (2), Solidago nemoralis (3), Sporobolus heterolepis (3), Viola pedatifida (1), Zigadenus elegans (2), Zizia aptera (5).

I developed this experiment to address questions about the impact native parasitic plants have on plant community members. In late October I harvested biomass from my pipp places to understand how species diversity and abundance change after planting parasites.

Start year: 2019

Location: Douglas County, Minnesota; exPt 1

Overlaps with: Experimental plot management, Hesperostipa common garden experiment

Materials collected: 216 .1 x 1m strips of dried biomass are stored at the Chicago Botanic Garden.

Data collected: Find data related to this project including the planting scheme in the cgdata repository in ~cgdata\summer2019\Hemiparasites (note/the key for HemiparaMap: C. umbellata = Blue, P. canadensis = Red, C + P = Purple, Soil plugs = Brown, Just seeds = Green).

2020 update: Aphid addition and exclusion

Team Echinacea continued the aphid addition and exclusion experiment started in 2011 by Katherine Muller. The original experiment included 100 plants selected from exPt01 which were each assigned to have aphids either added or excluded through multiple years. The intention is to assess the impact of the specialist herbivore Aphis echinaceae on Echinacea fitness.

The 2020 aphid team was Anna Allen and Allie Radin. They located 25 living exclusion plants and 16 living addition plants. The experiment was conducted from July 6th to August 19th, with the final visit consisting only of observation. Aphids were moved only during four visits from late July to mid-August due to late arrival and low numbers of aphids. Only one or two aphids were applied to each plant during each visit. They recorded the number of aphids present in classes of 0, 1, 2-10, 11-80, and >80. They also recorded the number of aphids added.

Aphids on an Echinacea leaf

Start year: 2011
Location: Experimental Plot 1
Overlaps with: Phenology and fitness in P1
Data collected: Scanned datasheets are located at ~Dropbox\teamEchinacea2020\allisonRadin\aphidAddEx2020.


  • 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

You can read more about the aphid addition and exclusion experiment, as well as links to prior flog entries mentioning the experiment, on the background page for this experiment.

2019 Update: Pollen addition and exclusion

Supplemental pollen — pollen that an Echinacea head might not otherwise receive—could increase a plant’s fitness. But does this extra pollination lead to a tradeoff in survival or flowering consistency? Since 2012, we have been manipulating the amount of pollen Echinacea plants receive – either no pollen, or lots of pollen – and recording how this affects their fitness and survival. In 2012 and 2013 we identified flowering E. angustifolia plants in experimental plot 1 and randomly assigned one of two treatments to each: pollen addition or pollen exclusion. The team bagged the heads of all plants and hand-pollinated the addition treatment, and did not manipulate the exclusion plants further. Plants receive the same treatment across years.

In summer 2018, 14 of the 26 plants alive in the pollen addition and exclusion experiment flowered, producing a total of 25 heads. This year none of those plants flowered. Of the original 38 plants in this experiment, 12 of the exclusion plants and 14 of the pollen addition plants are still alive. No plants died between 2018 and 2019. This year’s data were unique among the eight years of data collected, because not a single plant in the experiment produced even a single head. The dramatic decrease in flowering rates this year may help or hinder us in analyzing this data set and providing answers to this eight-year question.

Tris did not find significant demographic differences between plants which received pollen exclusion, addition or open pollination treatments.

Start year: 2012

Location: exPt1

Physical specimens: We harvested no specimens this year

Data collected: Plants survival and measurements were recorded as part of our annual surveys in P1 and can be found with the rest of the P1 data in the R package EchinaceaLab.

Michael presented a poster on the polLim experiment at MEEC 2019, which you can find here

Tris also presented a poster on polLim at MEEC 2019, which you can find here

You can find more information about the pollen addition and exclusion experiment and links to previous flog posts regarding this experiment at the background page for the experiment.

2019 Update: Heritability of fitness – qGen2 and qGen3

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)

Location: exPt8

Overlaps with: qGen1, 1996 and 1997 gardens, heritability of flowering time, common garden experiment, flowering phenology in experimental plots

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).

2019 Update: Inbreeding experiment – Inb2

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!

Summary of survival in inb2 by parental site.

For more summary plots, click these links:

Start Year: 2005 (crosses) and 2006 (planting)

Location: exPt1

Overlaps with: inb1, 1996 and 1997, common garden experiment, flowering phenology in experimental plots

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).


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

2019 Update: Reproductive Fitness in Remnants

Monitoring reproductive fitness in the remnant populations is a staple of Team Echinacea’s summer activities. Understanding the reproductive success of plants in remnant populations provides insight to a vital demographic rate contributing to the persistence (or decline) of remnant populations in fragmented environments.

In summer 2019, we harvested 40 seedheads to study patterns of reproductive fitness in 8 remnant Echinacea populations (ALF, EELR, KJ, NWLF, GC, NGC, SGC, NNWLF) (the same populations used where I studied phenology and gene flow). I randomly selected 1/3 of flowering heads at each remnant to harvest. In addition, I collected all seedheads from especially small or isolated remnants (specifically, GC, KJ, and the cluster of plants just north of EELR).

In early January, I dissected the seedheads. I extracted the achenes by row so that I will be able to observe temporal variation in seed set within heads. Ideally, next I will x-ray the achenes and assess seed set by observing the proportion of achenes that contain embryos. However, the x-ray machine at the Chicago Botanic Garden is currently out of service, so instead I may need to weigh or germinate the achenes to see if viable embryos are inside.

Extracting achenes by row, so that I know which achenes resulted from florets that flowered early (i.e., at the bottom of the seedhead) or late (i.e., at the top of the seedhead). Tedious but possible!

Start year: 1996

Location: Roadsides, railroad rights of way, and nature preserves in and around Solem Township, MN

Overlaps with: Phenology in the RemnantsGene Flow in Remnants

Products: We will compile seed set data from 2019 into a dataset with seed set data from previous years, which is located here:

You can read more about reproductive fitness in remnants, as well as links to prior flog entries mentioning the experiment, on the background page for this experiment.