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2020 Update: common garden experiments

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 (inb1inb2), and assessing quantitative genetic variation (qgen1). There are also a number of smaller experiments in it, including fitness of Hesperostipa sparteaaphid 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.

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).

Survival in common gardens

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!

2017 update: Heritability of fitness–qGen2 & qGen3

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.

2016 update: Heritability of fitness–qGen2 & qGen3

Alex and Lea measure qGen2 seedlings with much enthusiasm.

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.

The Flowers Are Coming

Today was another great day on Team Echinacea. This morning we continued our work on Q2, and we continued to make significant progress. We measured many plants and found at least five new plants in the experimental site. After a hearty lunch and a short time marveling at ice formations in a water bottle, Amy Dykstra gave a presentation on her research, which included her study of local adaptation of Echinacea. The afternoon was filled with preparations for IS projects (for the Wooster folks) and independent projects for the rest of us. Leah and Laura quickly became adept at catching pollinators, but were not so successful at transferring the collected pollen to fuchsin jelly. The rest of us hunkered down on our computers, read some literature, prepared project proposals and thought about how hard it would be to use NMR and IR to analyze pollen.

20160622_160359

Leah regales us with stories of captured pollinators and attempts at melting fuchsin jelly on a car dashboard.

The exciting news of the day is that we found our first flowering Echinacea in P1 and at Elk Lake Road East! Tomorrow we will find out how many more Echinacea are flowering.

The second flowering Echinacea (found at Elk Lake Road East)!

 

Must destroy all sweet clover!

The gang had a busy day today, almost all of it in the warm June sun. Alyson continued setting up her experimental plots in the Staffanson bog, and spent the afternoon measuring canopy cover and soil pH for her IS project. Meanwhile, the rest of the team (minus Gretel, who was setting up work for q2 juvenile counts) picked up our fleg begs and counted Hesperostipa spartea and weeded in p1. Amy and James found one H. spartea specimen with 137 seeds! We are now up to 17 out of 59 rows surveyed. Meanwhile, Will, Alex and Per led the crew in weeding out the non-native yellow sweet clover (Melilotus) from the periphery of the plot area. Hopefully we eliminated a lot of potential seeds form the seed bank, meaning that in future years the rows will be devoid of this weedy legume and the study Echinacea will have less competition. Stuart also showed me what poison ivy looks like for the third time, and I still don’t think I could pick it out of a lineup.

Per with a bundle of sweet clover picked from p1.

Per with a bundle of sweet clover picked from around p1. This is probably less than 10% of what was removed today.

After some brief (or for Alex, who was cleaning the bathroom, not so brief) chores at the Hjelm House, the team returned to exPt8 (experimental plot 8) to search for juvenile Echinacea  crosses planted in 2013. This meant more time bent over, although now instead of looking for seedlings we were looking for melted bits of toothpick (which were placed to mark seedlings). Some seedlings were in great shape — Alex and I found a couple with basal leaves over 10 cm tall. Others were not in great shape, either dead or missing like Jimmy Hoffa. We got about a third of the work for qGen2 this afternoon. It may rain tomorrow, so we’re bracing for indoors-work and hiding our bicycles inside.

Using a pink sword to claim the new seedling (left) for Team Echinacea. We used cocktail swords to denote seedlings germinating this year from seeds planted in 2013.

Using a pink sword to claim the new seedling (left) for Team Echinacea. We used cocktail swords to denote seedlings germinating this year from achenes sown in 2013.

Project status update: Heritability of fitness–qGen2 & qGen3

Pollen vials from Landfill sire 403 used in qGen3 crossing 2015

Pollen vials from sire 403 used in qGen3

The goal of the qGen2 and qGen3 experiments is to compare the evolutionary potential of two remnant prairie populations of Echinacea angustifolia by estimating the additive genetic variance of fitness under two mating scenarios: crosses performed within the core sites (core x core) and crosses performed between the core site and nearby sites (core x periphery). Additionally, we will test for differentiation among families; do progeny from sires differ after accounting for maternal (dam) effects?

In June 2015 we assessed survival and measured 1-year-old plants from qGen2. During the summer and fall of 2015 we replicated the qGen2 experiment through a second crossing experiment. We sowed the resulting achenes in exPt 8 as our qGen3 cohort.

Read more about the qgen2 and qgen3 experiments.

IMG_5251

Dam to be crossed with pollen from 2 sires

Start year qGen2: 2013

Start year qGen3: 2015

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

 

Summer fieldwork begins in Minnesota

The summer field season is off to a great start! We have assembled an excellent team to investigate ecology and evolution in fragmented prairie habitat focusing on the narrow-leaved purple coneflower as a model organism. Meet members of the team.

Team Echinacea 2015: Danny, Matt, Ben, Will,

Team Echinacea 2015: Danny, Matt, Ben, Will, Gina, Taylor, Lea, Amy, Katherine, Alison, Abby

We started the season with tours of local prairies large and small, including Staffanson Prairie Preserve, Hegg Lake WMA, which are large and protected. Stay tuned for team-members’ first impressions of some of the nearby remnant Echinacea populations.

Team-members hail from near & far: Barrett, Elbow Lake, and Alexandria, Minnesota & California, Alabama, New York, and Rhode Island. They are excited to develop summer projects and they will post their proposals here next week. Our team includes four college students, four who just graduated college, two high school teachers, and one high school student. And there are the old-timers.

To get ready for field work, we took the Hjelm House out of winter storage and cleaned out our storage facilities (g3). We inventoried supplies and made signs and tags for fieldwork. Everyone got a pouch with tools and supplies and Gretel has assigned us all a data collector. This may be (should be) the last year for our trusty handspring visor data collectors. The visors are trustworthy, but the computers and software that run them are showing their age.

The first main activity of the season was assessing survival and growth of 2526 plants in the Q2 experiment, which is designed to quantify the additive genetic variation in two Echinacea populations. The amount of additive genetic variation determines a population’s capacity for adaptation by natural selection. Genetic variation is very important for the persistence of populations in prairie habitat. We’ll find out how much variation Echinacea has, which will give us some ideas about future prospects for these populations in the rough-and-tumble and rapidly changing world out here.

We got rained out several times this week, but managed to measure all 2526 plants. We found a few plants that escaped detection last summer and we even found one seedling. Welcome to the experiment, fellas! We’ve got our eyes on you.

Overwinter survival appears to be quite good and most of the toothpicks we used to identify individual plants made it through the winter too. The tallest plants were just over 20 cm and some plants had 3 or more leaves. This is great news for plants that were sown as seed in fall 2013. Growth conditions are challenging: a cold winter with little snow, a dry spring, shading out by established plants, chewing by herbivores, … it’s a tough life for a prairie plant.

All in all, it promises to be a great summer. We’ll keep you posted.

broadcast seeding

Dwight and Stuart broadcast native prairie seed in experimental plots p1 & p8 on Friday. At 34 °F (1°C) it was the warmest day in a month. It was also very windy –great for spreading seed! We broadcast Bouteloua curtipedula, Schizachyrium scoparium, Galium boreale, and Phlox pilosa directly on the snow. There wasn’t much snow and it was melting. We broadcast Lathyrus venosus in p1. We stored about half of each species, except L. venosus, in the Hjelm house to broadcast in the spring. (Hedging our bets.)