2018 update: Demographic census in remnants

As always, demo was a huge job this year. We made some small adjustments to how we do total demo and flowering demo (we do flowering demo in every remnant now). Now, in our big plots, we only do a sample of the plants instead of total demo. Check out the updated table below (remember we did flowering demo everywhere). We still managed to shoot thousands of points, take even more demography records, and, as always, follow the colored flags!

Total Demo BTG, Common Garden,  DOG, East of Town Hall, KJs, Krusemark, Landfill West, Loeffler Corner East, Near Town Hall, Nessman, NNWLF, NRRX,   recruit he, recruit hp, recruit hs, recruit el, recruit hw, recruit ke, recruit kw, RRX,  RRXDC, South of Golf Course, Steven’s Approach, transplant plot, Tower, Town Hall, West of Aanenson, Woody’s
Sample Demo Aanenson, Around LF, East Riley, Hegg Lake, Elk Lake Road East, Golf Course, Landfill, Loeffler Corner, North of Golf Course, NWLF, On 27,  Riley, Staffanson,  Yellow Orchid Hill

Evan and Will doing demo by the corn. While you might not be able to see the echinacea, we promise, it’s under there!

So how do we do demo? When we find a new flowering Echinacea plant, we give it a tag and get its location with a survey-grade GPS (better than 6 cm precision). Then, we can revisit this plant for years to come and monitor its survival and reproduction. We’re monitoring over 10,000 plants as of 2018.

This season we added 3868 demo records and about 1539 survey records to our database. After over 20 years of this method, we are starting to get a very good idea of the demography of echinacea in Solem Township. With this many records, organizing demap in the future is going to be a big task!

Year started: 1996

Location: Roadsides, railroad rights of way, and nature preserves in and near Solem Township, Minnesota.

Overlaps with: Flowering phenology in remnants, fire and flowering at SPP

Data collected: demo records include Flowering status, number of rosettes, number of heads, neighbors within a 12 cm radius of plants found. These are all taken with PDAs that sync with an MS Access database. They are all transferred to the demap R repository in bitbucket with git version control.

GPS points shot: Points for each flowering plant this year shot mostly in SURV records, stored in surv.csv. Each location should be either associated with a loc from prior years or a point shot this year.


  • Amy Dykstra’s dissertation included matrix projection modeling using demographic data
  • Project “demap” merges phenological, spatial and demographic data for remnant plants

You can find out more about the demographic census in the remnants and links to previous posts regarding it on the background page for this experiment.

2018 update: Fire and flowering in SPP

For her REU project, Brigid gathered data to study the relationship between flowering density and seed set. She worked at Staffanson Prairie Preserve, which appears to have higher flowering density in burn years than non-burn years. This year, 2018, was a burn year on the east side of the preserve. Brigid and Team Echinacea kept track of the style persistence of about ~150 individuals many of  which we have phenology and style persistence information from prior years. These individuals were harvested and their achene count and seed set will be assessed by volunteers  and  interns at the CBG.

Brigid also observed nearest neighbors for many of the plants that she tracked. It might be the cases that echinacea flowers are more successful if they have other flowering plants nearby. Synchrony is a large part of why fire is so important, and, since SPP is our largest remnant prairie, it’s the best place to test the relationship between fire and synchrony. Number of heads, phenology, and head size may also \ interact with fire — we’ll know once we look at the data!

Site: Staffanson Prairie Preserve

Start year: 1996

Overlaps with: Phenology in Remnants, Reproductive Fitness in Remnants

Data and Samples: We shot 90 GPS points for nearest neighbors, many of which were plants that flowered for the first time this year. We also harvested 22 heads that are awaiting cleaning at CBG

Products: None so far

2018 update: Common garden experiment–1996 cohort

In 2018 only 19% of the plants flowered, despite it being a burn year. Is the 1996 finally showing its age?

In 2018, 51 plants flowered of the surviving 269 plants in the 1996 cohort. That means that 41% of the original plants are surviving and 19% of the living individuals flowered. That’s up huge since last year, where only 2% flowered, and the year before where five percent of living individuals flowered. In contrast, however, 45% of living plants flowered in 2015, and  37%, 34%, and 40% flowered in 2014, 2013, and 2012 respectively. We found that of the original 646 individuals, 269 were alive in 2018, only 15 fewer than last year. We are not sure why so many more plants flowered this year. It’s probable that the fire in the plot in fall 2017 influenced flowering rates.

The 1996 cohort has the oldest Echinacea plants in experimental plot 1; they are 22 years old. They are part of a common garden experiment designed to study differences in fitness and life history characteristics among remnant populations. Every year, members of Team Echinacea assess survival and measure plant growth and fitness traits including plant status (i.e. if it is flowering or basal), plant height, leaf count, and number of flowering heads. We harvest all flowering heads in the fall, count all achenes, and estimate seed set for each head in the lab. As yet, these heads are still waiting to be cleaned April 2019.

Start year: 1996

Location: Experimental plot 1

Overlaps with: phenology in experimental plots, qGen3, pollen addition/exclusion

Physical specimens:

  • We harvested 59 heads. At present, they await processing in the lab to find their achene count and seed set.

Data collected:

  • We used Visors to collect plant growth and fitness traits—plant status, height, leaf count, number of flowering heads, presence of insects—these data have been added to the database
  • We used Visors to collect flowering phenology data—start and end date of flowering for all individual heads—which is ready to be added to the exPt1 phenology dataset
  • Eventually, we will have achene count and seed set data for all flowering plants (stay tuned)


  • See the exPt1 core dataset where yrPlanted == ‘1996’ for 1996 cohort fitness measurements

You can find more information about the 1996 cohort and links to previous flog posts regarding this experiment at the background page for the experiment.

2018 update: Richardson’s Liatris and Solidago phenology

In the summer or 2018, Lea collected data for the third year of her observational study quantifying

Flowering Liatris

flowering phenology and reproductive success (seed set) for Liatris aspera and Solidago speciosa plants located along a transect at Staffanson Prairie Preserve. Staffanson is divided into east and west units. The west unit of Staffanson was burned Spring 2016. In 2016, Lea looked for differences in phenology and reproduction of east vs. west Liatris and Solidago plants. In 2017, neither unit was burned. In 2018, the east unit burned. Data collected this year combined with data collected in 2016 and 2017 will enable us to to see if burns influence phenology or reproduction. To assess phenology, Lea visited plants three times a week and recorded if they were flowering. She took GPS data for each plant included in the study. She also mapped the seven nearest neighbors of all flowering plants within her transect in 2018. Additionally, Lea visited all plants in the 2016 and 2017 datasets to see if they were still present and if they were flowering. To assess reproduction (seed set), plants were harvested and brought back to the Chicago Botanic Garden so that seeds could be removed from the plant and x-rayed. This study helps us understand how fire, phenology, and reproduction are linked in species that are related to Echinacea angustifolia.


Start year: 2016

Location: Staffanson Prairie Preserve

Overlaps with: Fire and fitness of EA, Flowering phenology in remnants

Physical specimens: 

  • ~80 harvested Liatris aspera specimens from summer 2018, located at the CBG
  • ~80 harvested Solidago speciosa specimens from summer 2018, located at the CBG

Data collected: Phenology data was taken on the visors every Monday, Wednesday, and Friday through the growing season. Paper harvest data sheets were used and brought back to CBG.

GPS points shot: ~543 GPS points were visited or shot, one point was visited for each plant monitored in summer 2016 and 2017, and points were shot for each plant in the 2018 dataset along with its seven nearest neighbors.

2018 update: Pollen addition and exclusion

Michael adds pollen to one of the “addition” plants using a toothpick and a vial of collected pollen.

Does receiving the maximum amount of pollination vs. no pollen at all affect a plant’s longevity or likelihood of flowering in subsequent years? Since 2012, we have been either adding or excluding pollen from the same plants’ heads to test what role pollen plays in ongoing survival and flowering. Unlike the previous two years, many plants in this experiment flowered this year, and analyzing them will likely reveal interesting results.

This summer, 14 of the 26 plants remaining in the pollen addition and exclusion experiment flowered, for a total of 25 heads. This is the most heads the pollen addition and exclusion project has had since 2013. We continued experimental treatments on these flowering plants and recorded the survival and flowering status of all plants in the experiment. Of the original 38 plants in this experiment, 12 of the exclusion plants and 14 of the pollen addition plants are still alive.

In this experiment, we assess the long-term effects of pollen addition and exclusion on plant fitness. 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. When plants flower in subsequent years they receive the same treatment they were originally assigned. The increase in flowering rates this year will allow us to move forward with analyzing this data set, to potentially answer this 7-year question.

Start year: 2012

Location: Experimental plot 1

Physical specimens: We harvested 25 flowering heads from this experiment that will be pulled from the normal ACE processing and pushed through faster. This is so we can more quickly determine achene count and proportion of full achenes.

Data collected: Plants survival and flowering data was recorded as part of our annual surveys in P1 and can be found with the rest of the P1 data. Data on when heads had pollen added can be found in CGData/115_pollenLimitation, particularly dailyPollenAddition for the year 2018.

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.

2018 update: Aphid addition and exclusion

This summer Team Echinacea continued adding and excluding aphids to plants in the experiment that Katherine Muller started in 2011. Katherine Muller randomly designated a sample of 100 Echinacea plants in experimental plot 1 for either aphid addition or removal.

Andy managed the project for the team in 2018, making sure that aphids were removed from the exclusion plants, and added to the addition plants. Twice a week, Andy (and occasionally Morgan) visited every plant in the study, recording the number of aphids, ants, and leaves infested. There remain 54 plants in the aphid study, 26 from the aphid addition plants and 28 from the exclusion plants. The data for this year will be added to the ongoing dataset.

Aphis echinaceae is a specialist aphid that is found only on Echinacea angustifoliaRead more about this experiment.


That’s a lot of aphids!

Start year: 2011

Location: Experimental Plot 1

Overlaps with: Phenology and fitness in P1

Data collected: All sheets describing the addition and removal of aphids from echinacea are stored in two places. All physical sheets are in Stuart’s manila folder titled “Aphid Add/Ex 2018.” Additionally, all data from the sheets are present in the aphidAdd2018Master and aphidEx2018Master. The exact path of this file may change but is currently ~\Dropbox\teamEchinacea2018\andyHoyt\aphidAddEx2018\aphidDataAllYears

Physical specimens: We harvested 25 flowering heads from this experiment that will be pulled from the normal ACE processing and pushed through faster. This is so we can more quickly determine achene counts and proportions of full achenes.


  • 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 previous flog entries mentioning the experiment, on the background page for this experiment.

2018 Update: Pollen to Seed

Over 90% of flowering plants rely on insect pollinators for sexual reproduction. Pollination biologists often quantify pollinator effectiveness by counting the number of pollen grains a pollinator deposits on a plant’s stigma in a single visit. However, flowers in the Asteraceae, like Echinacea, are uniovulate, meaning that a there is just one ovule per style. Therefore, it theoretically only takes one viable pollen grain to fully set seed in an Echinacea ovary, which means the standard method of quantifying pollinator effectiveness may not be appropriate for Echinacea and other members of the Asteraceae. In this study, we wanted to estimate how many pollen grains are really necessary to set a seed in Echinacea angustifolia. We performed 60 hand crosses with varying levels of pollen deposited. Each cross consisted of nine total styles of which three were removed before the cross to get estimates of self-pollen deposition, and six styles were crossed with pollen from other flowering individuals. We then collected styles and stained them using fuchsin gel and to count the number of pollen grains using a compound microscope. We x-rayed the achenes, fruits, from the crosses to determine if a seed was present. Preliminary data shows that seed set varies with the pollen grains deposited. These data will advance our understanding of pollinator effectiveness in this uniovulate plant. By increasing our understanding of pollinator efficiency in the Asteraceae we can better predict the consequences of pollinator declines in fragmented habitats, such as the North American tallgrass prairie.

Mia doing one of her many, many crosses

Start year: 2018

Location: exPt2

Overlaps with: The Big Event, Bees Remove and Deposit Pollen

Physical specimens: 183 Echinacea heads were dissected at the College of Wooster in fall 2018. They are currently being processed at CBG.

Data collected: Contact Dr. Jennifer Ison or Mia Stevens for data related to this experiment

Team members who have worked on this project include: Mia Stevens (2018) is the lead on this project.

2018 Update: Pollinators influencing male and female fitness a.k.a The Big Event

In plant populations where reproduction is mate-limited, the potential exists for selection on floral traits through both male and female function (seed production and siring success). Selection could be strong because of high variation in both male and female fitness. Additionally, most plants rely on a number of generalist insect pollinators, each of which is likely imposing selection on plant traits associated with reproduction. In many plant species, these generalist pollinators are native solitary bees. However, pollination research mainly focuses on large social bees—bumblebees and the non-native honeybee.

The main objective of this research project is to quantify how four generalist solitary bee taxa contribute to male and female fitness in the mate-limited prairie perennial, Echinacea angustifolia. To accomplish this objective, 15 researchers worked together to monitor the complete pollinator visitation in a prairie community with over 200 flowering Echinacea individuals during five days in July 2018. Through this effort, we recorded over 700 individual bee visits to a flowering Echinacea plant. To assess seed set, we removed achenes from Echinacea seedheads produced during the five observations day and xrayed the achenes for viable embryos.

Currently, we are germinating the resulting seeds and taking leaf samples. This summer, we will extract DNA from the leaf samples and genotype each sample at 11 previously optimized Echinacea-specific microsatellite loci. We will use a full paternity maximum likelihood analysis to quantify siring success mediated by the different generalist bee taxa. Our results will advance our understanding of relationships between male, female, and total fitness in plant populations. Our results will also reveal how different pollinators may cause variation in fitness. These results can help us predict the effects of changing pollinator communities for fragmented plant populations.

Everyone in exPt2 looking for bees

Start year: 2018

Location: exPt2

Overlaps with: Bees Remove and Deposit Pollen, Pollen to Seed

Physical specimens: 183 Echinacea Heads are currently being processed at CBG

Data collected: Contact Dr. Jennifer Ison for data related to this experiment

Team members who have worked on this project include: Everyone in 2018! The big event was just that- a few very big events

2018 Update: Reproductive Fitness in Remnants

In summer 2018, I harvested 80 seedheads from 12 remnant Echinacea populations (ALF, EELR, KJ, NWLF, GC, NGC, SGC, NNWLF, LC, RRX, NRRX, YOH) to study patterns of reproductive fitness. I sampled heads in two ways – (1) I randomly selected 20% of the individuals at each site (43 individuals) and (2) I randomly sampled up to 5 individuals from full factorial combinations of high, medium, and low spatial isolation and early, peak, and late flowering time (i.e., high spatial isolation/early flowering, high spatial isolation/peak flowering, etc.) across all sites (37 individuals).

In January 2019, I dissected seedheads that I collected from the NW sites (ALF, EELR, KJ, NWLF, GC, SGC, NGC, KJ, NNWLF). I extracted the achenes by row to observe temporal variation in seed set within heads. I x-rayed the achenes and assessed seed set in January.

Xray images that show whether achenes contain embryos or not

Start year: 1996

Location: Remnant prairies in central Minnesota

Overlaps with: Phenology in the Remnants, Gene Flow in Remnants

Products: Check back with the flog for preliminary results and annual reports.

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.

2018 Update: Gene Flow in Remnants

In summer 2018, I began a project to look at pollen movement within and among the remnant populations. To do this, I chose two focal areas, the NW sites in the study area (populations: ALF, EELR, KJ, NWLF, GC, SGC, NGC, KJ, NNWLF) and the SW sites (populations: LC, NRRX, RRX, YOH, and two large populations in between these sites). I mapped and collected leaf tissue from all individuals in the study areas and harvested seedheads from a subset of these individuals (see Reproductive Fitness in Remnants). I am currently extracting the DNA from the leaf tissue samples and a subset of the seeds I collected, and will use the microsatellite markers that Jennifer Ison developed in her dissertation to match up the genotypes of the offspring (i.e., the seeds) with their most likely father (i.e., the pollen source).

An Echinacea that has had today’s load of pollen fully removed by pollinators

Start year: 2018

Location: Remnant prairies in central Minnesota

Overlaps with: Reproductive Fitness in Remnants, Phenology in the Remnants

Products: Check back with the flog for preliminary results and annual reports.

Team members who worked on this project include: Amy Waananen