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 the phenological data collected this summer, we explore how flowering phenology influences reproductive fitness and estimate the heritability of flowering time in E. angustifolia.
In summer 2021, we visited 2,010 positions of the 3,961 positions originally planted. We measured 1,591 living plants, of which 681 were flowering, with a total of 1,283 flowering heads. In the fall, we harvested ~615 heads from exPt02. We do not have an exact number of heads harvested from exPt02 yet because we have not had time to complete head inventory. 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.
Measuring p2 involved 9 different people working a total of 4295 minutes (71.58 hours) on 4 separate days.
Physical specimens: We harvested approximately 615 heads from exPt 2. Many heads were harvested by ground squirrels, so our number of heads does not match the number harvested. We brought the harvested heads back to the lab, where we will count fruits and assess seed set for each head.
Data collected: We collected data on each position planted. We recorded whether the plant was alive; if it was alive, whether it was flowering or basal. Basal plants had information about number of basal rosettes, number of basal leaves, and length of longest leaf. For flowering plants, we collected rosette counts and the height of each flowering head. When harvesting, we collected samples of loose achenes and recorded where they were from. Next spring, we will search for seedlings at these positions to ensure that there are not any rouge plants.
Products: Will and Jennifer are working on a manuscript using the flowering phenology data from this plot.
You can find more information about the heritability of flowering time and links to previous flog posts at the background page for the experiment.
In 2017, we monitored the start and end dates of flowering for the 676 flowering plants (1116 heads) in experimental Plot 2. The first head started shedding pollen on June 26 and the latest bloomer ended flowering on August 19. Peak flowering was on July 13th. Note that these dates are subject to change as this is preliminary data that has not been fully cleaned and analyzed.
To examine the role flowering phenology plays in the reproduction of Echinacea angustifolia, Jennifer Ison planted this plot in 2006 with 3961 individuals selected for extreme (early or late) flowering timing, or phenology. Using the phenological data collected this summer, we will explore how flowering phenology influences reproductive fitness and estimate the heritability of flowering time in Echinacea angustifolia.
One of the earlier flowering plants at exP2 this summer, a plant with 5 heads at Row 2 Position 1.
Start year: 2006
Location: Experimental Plot 2, Hegg Lake WMA
Overlaps with: phenology in experimental plots, phenology in the remnants
Physical specimens: We harvested approximately 1081 heads from exPt 2 (preliminary inventory). Some of these heads had a major loss of achenes, either due to the early flowering time that we were not expecting or windy and rainy weather that dispersed the achenes quickly. We brought the harvested heads back to the lab, where we will count fruits and assess seed set for each head.
Data collected: We visited all plants with flowering heads every two days (three days after weekends) until they are done flowering to record start and end dates of flowering for all heads. We will manage phenology data in R and add it to the full dataset.
Products: Will estimated heritability of flowering time using data from 2015 and presented his findings last summer at ESA (see his poster). He is continuing this work by assessing how heritability estimates differ between years in burn and non-burn years, now including 2016.
You can find more information about the heritability of flowering time and links to previous flog posts at the background page for the experiment.
To examine the role flowering phenology plays in the reproduction of Echinacea angustifolia, Jennifer Ison planted experimental plot 2 (exPt 2) in 2006 with 3961 individuals selected for extreme (early or late) flowering timing, or phenology. In 2016, we monitored the start and end dates of flowering for the 570 flowering plants (933 heads) in the plot. The first head started shedding pollen on June 22 and the latest bloomer ended flowering on August 8th. Peak flowering was on July 7th, when 810 heads were flowering. Using the phenological data collected this summer, we will explore how flowering phenology influences reproductive fitness and estimate the heritability of flowering time in Echinacea angustifolia.
Tracking phenology for 900+ heads in exPt2 was a big job. Here, three teams assess phenology on a nice day at exPt 2.
Start year: 2006
Location: Experimental Plot 2, Hegg Lake WMA
Overlaps with: phenology in experimental plots, phenology in the remnants
Physical specimen: We harvested 870 heads from exPt 2. We were unable to harvest some heads which had been grazed by rodents. We brought the harvest back to the lab, where we will count fruits and assess seed set. Jennifer previously collected tissue samples from all individuals in the plot and plans to use these to genotype all of the individuals that flowered in 2016 this year and determine their parentage in exPt 1.
Data collected: We visit all plants with flowering heads every three days until they are done flowering to record start and end dates of flowering for all heads. We managed phenology data in R and added it to the full dataset.
Products: Will estimated heritability of flowering time using the data from 2015 and presented his findings this summer at ESA (see his poster here). He is continuing this work by assessing how heritability estimates differ between two years. He is comparing flowering in 2015 (a burn year) and 2016 (a non-burn year).
You can find more information about our experiment the heritability of flowering time and links to previous flog posts regarding this experiment at the background page for the experiment.
Description: This experiment is designed to study the heritability of flowering time and to examine the role flowering phenology plays in the reproductive fitness of Echinacea.In 2005, Jennifer Ison and Team Echinacea assessed phenology in experimental plot 1 and identified individuals with extreme (i.e. early or late) flowering time (Ison and Wagenius 2014). These extreme individuals served as maternal plants for this experiment; we harvested their achenes, which we then grew into seedlings in the lab. Jennifer collected tissue from the seedlings to identify the pollen donor (i.e. the paternal plant) with microsatellite markers she developed (Ison et al. 2013). In 2006, Jennifer and other team members planted the seedlings (3961 in total) in experimental plot 2. Each year since they began flowering (1 plant flowered in 2009), team members have assessed flowering phenology of the individuals in experimental plot 2.
Experimental plot 2
We identify flowering plants early in the season, before they begin shedding pollen, and assess their flowering status regularly throughout the season until they are done flowering (no more than 3 days between observations). In doing so, we collect data on start and end date of flowering for all individual heads in the plot.
To learn more about the beginnings of this large seedling growth experiment read this description.
Start year: 2006
Location: Hegg Lake WMA; experimental plot 2
Products:
These three papers are about the flowering phenology and mating patterns of the parents of the plants in experimental plot 2:
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
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
Ison, J.L., S. Wagenius, D. Reitz., M.V. Ashley. 2013. Development and evaluation of microsatellite markers for a native prairie perennial, Echinacea angustifolia (Asteraceae). Applications in Plant Sciences 1: 1300049. PDF
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).
Each year, we assess flowering phenology in experimental plots to determine mating potential for individual plants and see how a number of factors may affect flowering phenology. Some of the factors we have investigated in the past include heritability, burning, and climate.
2019 was truly a special year for Echinacea flowering phenology in experimental plots. There were flowering plants in four – yes FOUR – experimental plots. We had the usual flowering plants in exPt1 and exPt2 at Hegg Lake. We also had a flowering plant in exPt8 (qgen2 and qgen3) and exPt9 at Hegg Lake. Unfortunately, we did not see the flowering plant with an E. pallida dam at exPt9 until late in the season, so we did not take phenology in exPt9.
This Echinacea head is mid-flowering. It has more than 2 rows shedding pollen and more than 11 immature florets.
This year, we visited the three other plots and followed the usual Echinacea phenology protocol. We recorded first flowering day and subsequently recorded dates of “mid” and “late” flowering. Finally, we recorded the final flowering date of each plant.
In addition to the single flowering plants in exPt8 and exPt9, exPt1 had 63 flowering heads we tracked for phenology and exPt2 had a whopping 1207! The first flowering head in exPt1 started on July 3rd, while the first head in exPt2 started flowering on July 1st. The last day of flowering in exPt1 and exPt2 was August 21st. What a long summer of taking phenology data!
Data/
materials collected:phenology data (start date, mid flowering, end date,
etc…), harvested heads for the ACE protocol. All phenology data can be found in
the cgData repository in the subfolder p1p2Phenology.
This year, the number of flowering plants in our main experimental plot (exPt1) dropped in half compared to last year. This might be due to the lack of a burn in the prior fall or spring. Plot 2 (exPt2) had about the same number of heads in ’16 & ’17.
In exPt1, we kept track of approximately 72 heads. The peak date was July 19th. The first head started flowering on July 2nd and the last head finished up on August 21st. In contrast, we kept track of 1076 heads in exPt2, about 140 more than last year! The peak date for these Echinacea was a bit earlier, July 13th. exPt2 heads also started and ended earlier (June 22 – August 19).
We harvested the heads at the end of the field season and brought them back to the lab, where we will count fruits (achenes) and assess seed set.
Flowering schedules for 2017 in exPt1 and exPt2. Black dots indicate the number of flowering heads on each date. Gray horizontal line segments represent the duration of each head’s flowering and are ordered by start date. The solid vertical line indicates peak flowering, while the dashed lines indicate the dates when 25% and 75% of heads had begun flowering, respectively. Note the difference in y-axes between the two plots. Click to enlarge!
Physical specimens: Harvested heads from both experimental plots are in the lab at CBG. The ACE protocol for these heads will begin soon.
Data collected: We visit all plants with flowering heads every 2-3 days starting before they flower until they are done flowering to record start and end dates of flowering for all heads. We managed phenology data in R and added it to our long-term dataset. The figures above were generated using package mateable in R. If you want to make figures like this one, download package mateable from CRAN!
You can find more information about phenology in experimental plots and links to previous flog posts regarding this experiment at the background page for the experiment.
Every year we keep track of flowering phenology in our main experimental plots, exPt1 and exPt2. Fewer plants than usual flowered in exPt1 in 2016: 149 plants (179 heads) flowered between June 24th and August 7th. The population’s mean start date of flowering was July 5th and the mean end date was July 18th. Peak flowering in 2016 was on July 10th, when 143 heads were in flower. For comparison, peak flowering in 2015 was on July 27th, when there were nearly 10x as many heads flowering as on this year’s peak. The earlier phenology and low numbers of flowering we observed this year relative to 2015 is likely due at least in part to the plot burn schedule (2015 was a burn year and 2016 was a non-burn year), but there were still many fewer flowering plants than any season, burn or non-burn, in the past 10 years.
We kept track of 934 flowering heads in ExPt2, where the first head started shedding pollen on June 22 and the latest bloomer ended flowering on August 8th. Peak flowering was on July 7th, when 810 heads were flowering. ExPt2 was designed to study the heritability of phenology—you can read more about progress of that experiment in the upcoming 2016 heritability of phenology project status update.
At the end of the season we harvested the heads and brought them back to the lab, where we will count fruits (achenes) and assess seed set.
ExPt1 and Expt2 flowering schedules from 2016. Dots represent the number of flowering heads on each date. Horizontal line segments represent the duration of each heads flowering and are ordered by start date. The solid vertical line indicates peak flowering, while the dashed lines indicate the dates when 25% and 75% of heads had begun flowering, respectively. Click to enlarge!
Physical specimens: We harvested 177 heads from exPt1 and 870 from exPt2. Attentive readers may note that we harvested about 64 fewer heads than we tracked for phenology. That’s because before we could harvest many seedheads at exPt2, rodents chewed through their stems and ate some fruits (achenes). We recovered most of the heads that were grazed from the ground and made estimates of number of fruits lost due to herbivory, but we couldn’t find some heads. Arg. We brought the harvest back to the lab, where we will count fruits and assess seed set.
Data collected: We visit all plants with flowering heads every three days until they are done flowering to record start and end dates of flowering for all heads. We managed phenology data in R and added it to the full dataset. The figure above was generated using package mateable in R. If you want to make figures like this one, download package mateable from CRAN!
You can find more information about phenology in experimental plots and links to previous flog posts regarding this experiment at the background page for the experiment.
Every year Team Echinacea monitors the timing of flowering, or phenology, of individuals in the experimental plots. Timing matters because both abiotic conditions, such as temperature and photoperiod, and biotic conditions, such as mate availability and pollinator abundance change over the course of a season and may have consequences for reproductive fitness. This study began in 2005 in experimental plot 1 when Jennifer Ison investigated how an individual plant’s timing within a season affected its mating opportunity and reproductive fitness (Ison and Wagenius 2014).
A plant is considered flowering when it produces pollen– this head looks like it is on its second day of flowering.
Each year, team members identify all plants that are going to flower that year before they begin shedding pollen. We then make regular observations, no more than 3 days apart, of every head on every plant until they are all done flowering. In doing so, we obtain both flowering start and end dates for all flowering heads. We also collect information on any unusual phenological occurrences such as premature end of flowering due to disease, herbivory, or other causes.
Start year: 2005
Location: Experimental Plot 1 (at the research base) and Experimental Plot 2 (at Hegg Lake WMA).
Products:
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
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
Overlaps with: heritability of phenology, common garden experiment, phenology in the remnants, phenology of Echinacea pallida
Link to flog posts: Read updates and annual reportsabout this experiment on our flog (field blog) as written by members of Team Echinacea.
For his REU research project, Will is investigating heritability of flowering phenology in experimental plot 2 (p2), which was planted in 2006 with 3961 individuals selected for extreme (early or late) flowering phenology. This summer Will and Team Echinacea monitored the phenology of all 646 flowering plants (1216 individual heads) in p2, as well as the phenology of their parents in p1. Flowering in p2 began on the 4th of July and ended on August 26th. The peak day of flowering was July 27th when 1018 heads were flowering. The average duration of flowering for a head was 12.1 days. It was a huge year for flowering in p2 with nearly 5 times more heads than 2014 and over 16 times more than 2013. Will is comparing the flowering schedules of the p2 plants with the 2005 phenology records of their parents. As the plants in p2 mature, and perhaps flower more frequently, we will continue to gather data on family lines of Echinacea to discern the genetic component of their flowering phenology.
Many flags indicate many flowering plants in experimental plot 2
Start year: 2006
Location: Experimental plot 2 (Hegg Lake WMA)
Overlaps with: phenology in experimental plots
Products: The 2015 phenology records from p2 will be added to the existing multi-year ExPt2 phenology dataset. Will is using tools from the R package echPhenology developed by Team Echinacea to analyze and produce visualizations of the flowering schedule observed in p2. He plans to present his findings at a conference in spring or summer 2016.
2015 flowering schedule of all heads in p2: First day of flowering was July 4th, peak was July 27th, and the last day was August 26th.
