APS Bulletin 398.Indd

The AMERICAN PEONY SOCIETY Bulletin

Photo courtesy Teppi Kinnari THE AMERICAN PEONY SOCIETY MEMBERSHIP & THE APS BULLETIN (APS) is a nonprofit horticultural orga- Dues are paid for the calendar year. nization incorporated as a member- Dues received before August 25 are re- ship corporation under the laws of the corded for the current year and mem- State of Missouri. APS is organized ex- bers will be sent all four issues of The clusively for educational and scientific Bulletin for that year (while supplies purposes, and especially to promote, last). Dues received between August encourage and foster the development 25 and November 25 will receive the and improvement of the genus Paeo- December issue of The Bulletin and nia and public interest therein. These all issues for the following year. Mem- purposes are expressly limited so that berships received after November 25 APS qualifies as an exempt organi- will be recorded for the following year. zation under section 501(c)(5) of the Online reading is available for the five Internal Revenue Code of 1954 or the most current Bulletin issues. Those corresponding provision of any future with online-only membership will not Internal Revenue law. Donors may not receive printed Bulletins. Membership deduct contributions to APS. information and an online registration Opinions expressed by contributors to form are available on the APS website. this publication are solely those of the Individual memberships are for one individual writers and do not necessar- or two persons at the same address, ily reflect the opinions of the APS Edi- receiving one copy of The Bulletin. tor, Officers, or Board of Directors. Commercial memberships are for one business and one or two persons at APS publishes paid advertisements as the same address. Refunds are avail- a service to its members, but cannot be able only to Institutional members for responsible for the quality of merchan- unmailed issues. All memberships are dise or services from these advertisers. non-transferable. Please notify APS promptly of address changes. Missed ISSN 0003-0384 issues must be claimed within one ©AMERICAN PEONY SOCIETY year. APS is not responsible for undeliv- All Rights Reserved erable mailing addresses.

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Summer 2021 Issue No. 398

COLUMNS President’s Message 4 Managing Soils Bulletin Board 5 Jim Sampson Page 7-13 DEPARTMENTS Nomenclature 6

2021 APS VIRTUAL CONVENTION Splendid Species If Wishes Were Peonies Page 16-17 Photo Contest & Virtual Exhibition 14-15

REGIONAL NEWS From Wasteland to Peony Park Duluth Peony Society 41 Leena Liljestrand Minnesota Peony Society 41 Page 21-24 Wisconsin Peony Society 42

The Language of Peonies A New “Unified” Plumule – Page 18 Classification for the Genus Paeonia

Part 1 – Donald R. Smith Page 25-40

PUBLISHED QUARTERLY IN SPRING, SUMMER, AUTUMN, AND WINTER President’s Message Summer 2021 Nate Bremer, APS President and Director – Reedsville, Wisconsin

What’s on my mind? Getting outside and working with peonies! At the time of this message (late March,) we are completing our “month of mud” which prohibits garden work as the ground thaws here in northeast Wis- consin. The woody peonies are beginning to break dormancy and there is much promise for a beautiful bloom season, but also reservation due to fickle weather patterns. In a short couple of weeks we’ll begin to see the herbaceous “nos- es” pop out of the ground and then the fields will have a color other than brown. I can’t wait! The past couple of months have been busy ones considering most of my time has been spent behind a computer readying for bloom season, an American Peony Society Virtual Convention offering and looking forward to the coming year’s APS events. The pandemic may have slowed the appearance of activity within APS; however, Society volunteers have been very busy. Membership has increased with more people looking to spend time in their gardens, keeping our Membership Chair and Treasurer extremely busy. The APS Seed Distribution Program has been running full speed with many orders with people looking to the future. The Peony Registry is being updated and saw more than 30,000 visits in Janu- ary. Conventions in Michigan and New York are being planned (they are looking really good). Our members are submitting articles and images for the Bulletin at record pace, which our Editor loves! Peony questions and Society inquiries through our Corresponding Secretary have seen break neck pace. New peony cultivar registrations continue to be submitted by hybridizers. Awards planning and nominations for the Gold Medal Peony and Personal recognition are being made by members and the Awards Committee is working to organize them for 2021. Many other activities and programs continue their quiet and behind the scene’s work – all pointing to a fantastic year ahead of us! As we move into the 2021 peony season I invite all members to become more involved in your society – I believe you will enjoy the camaraderie and find ful- fillment! All the Best, Nate Bremer

398-4 Bulletin Board 1) Since the in-person convention in Minnesota was cancelled this year due to the Covid-19 Pandemic, APS has been working on an exciting alternative/Virtual Convention for members. Details may be found on the APS website: https://americanpeonysociety.org/ news-events/if-peonies-were-wishes/.

2) The Peony Registry has been undergoing some work to improve details. When looking at the Registry you may see an icon like the one at right. This icon marks cultivars in which APS does not have an image for on file. Many are older cultivars which may no longer exist in commerce, but may reside in a member garden. APS is hoping our members may be able to assist with cultivars which are missing an image by donating a copy to the Registry. Images may be easily uploaded through the Membership Portal under the “Contribute” link at: https://americanpeonysociety.org/member-portal/pages/ contribute/submit-photos/.

3) The 2021 Seed Distribution Program Seed List is posted for Mem- bers interested in ordering and growing peonies from seed. Scott Parker, the SDP Program Chair, wishes to thank all donors for this popular APS program. Further details may be found on the APS website at: https://americanpeonysociety.org/seed-program/ seed-list/. As peony bloom season will be arriving shortly, please consider making some cross pollinations to share seeds with Membership through the SDP in 2022. The SDP welcomes all donations and it’s a great way to share your enthusiasm for peonies.

4) APS is aware some of our overseas members have experienced delays or difficulties with the delivery of The Bulletin. The Editor is investigating this issue with our publisher’s international mail house. The latest information we have is permit bulk mail (The Bulletin) is likely delayed due to pandemic impacts on postal services worldwide. Please be patient, as we believe Bulletins may arrive soon. If you are anxious to read the latest Bulletin, please remember it is available to all Members online through the Mem- ber Portal.

398-5 NOMENCLATURE Reiner Jakubowski, APS Registrar — Waterloo, Ontario, Canada The American Peony Society serves as the International Cultivar Registration Au- thority for Peonies and follows the rules set forth by the International Code of No- menclature for Cultivated Plants in the registration of new varieties.

Photographs and full descriptions can be found in the official PeonyRegistry, maintained on the American Peony Society Website at americanpeonysociety.org/cul tivars/peony-registry.

APS publishes a separate, Annual Registry Directory with parentage, full descriptions and photos, which will be mailed to all current members in late December. 'Spring Sun' [Herbaceous Hybrid] — Don Hollingsworth, Maryville MO, USA. 'Imperial Fuchsia' (Lactiflora Group) — Don Hollingsworth, Maryville, MO, USA. 'Sentry' [Herbaceous Hybrid] — Don Hollingsworth, Maryville, MO, USA. 'Enchanted' – [Herbaceous Hybrid] — Don Hollingsworth, Maryville, MO, USA. 'OSIRIS Mariette Pelland' (Lactiflora Group) — Serge Fafard, Saint-Thomas de Joliette, QC, Canada. 'Sandra L. Anderson' (Itoh Group) — Roger F. Anderson / David Cartwright, Whitewater, WI, USA. 'Schopenhauer' [Herbaceous Hybrid] — Roger Anderson / David Cartwright, Whitewater, WI USA. 'Rockin’ the Spirit' (Suffruticosa Group) — Roger F. Anderson / David Cart- wright, Whitewater, WI, USA. 'Good Karma' [Herbaceous Hybrid] — Roger F. Anderson / David Cartwright, Whitewater, WI, USA. 'Abbess Roothing' [Herbaceous Hybrid] — Harald W. Fawkner, Falköping, Sweden. 'Khuit' [Herbaceous Hybrid] — Harald W. Fawkner, Falköping, Sweden. 'Ossora' [Herbaceous Hybrid] — Harald W. Fawkner, Falköping, Sweden. 'Patience Spokoyny' [Herbaceous Hybrid] — Harald W. Fawkner, Falköping, Sweden.

398-6 What Works for Us

Jim Sampson - Mt. Hood, Oregon

For me, ‘dirt’ is the debris on my neither “dust to dust nor ashes trousers and under my nails, to ashes,” but in this case mostly while ‘SOIL’ is the living, breath- cycling as “gases to gases.” Con- ing, miraculous substance sup- sidering the sources it’s not sur- porting plant life itself. Its sci- prising that a cord of firewood entific management is deeply recycles almost entirely into car- challenged by the intricately bon monoxide, carbon dioxide, complex interactions of chemical, and water vapor (CO, CO2 & H2O). physical, biological, and environ- Easy come, easy go! It is the 14 mental factors. I am not a “soils” other elements that challenge us expert and do not presume such. as growers. Upon request I share the following thoughts on soil testing, fertil- Three are the familiar ‘Macronu- izing and management practices. trients’ nitrogen, phosphorous, Superb information and tutorials and potassium shown as the are available online from state N-P-K percentages on fertilizer extension services, fertilizer in- packages. Three more rank as the dustry consortiums, YouTube vid- ‘Secondary Nutrients’ calcium, eos, etc. Some gardening blogs magnesium, and sulfur (Ca, Mg, provide valuable advice, while S). Finally, there are eight ‘Micro- others seem to me questionable. nutrients’ (aka ‘trace elements’) including boron, chlorine, man- Seventeen essential elements are ganese, iron, nickel, copper, zinc, required for plant growth and re- and molybdenum (B, Cl, Mn, Fe, production. Three are fundamen- Ni, Cu, Zn, Mo). These rankings tal, free, mostly ubiquitous, and are based on greatest use by provided by air and water: Car- plants, not by most essential. bon, Hydrogen, Oxygen, (C-H-O). Alternatively, an excess supply, These eponymous essentials of especially for the micronutrients, carbohydrates include the sug- can easily be phytotoxic. ars, starches and cellulose being

398-7 Beyond nutrient needs, abiotic samples be taken? I rarely soil factors including soil tempera- sample here. My testing is limit- ture, texture, parent minerals, ed to ground that has been fal- water holding capacity and low pasture land, uncropped for drainage, gas exchange, organic several years and being used for matter, and pH (acidity/alkalin- peonies for the first time. My goal ity) play equally critical roles in is to establish the baseline of nu- the plant’s rhizosphere. trients present so as to amend the soil for the new peony plant- When applied to plants, Liebig’s ings. Initially I sampled both the Law of the Minimum states that top eight inch profile (0 to 8 inch “growth is restricted to by the depth) and the next eight inch- most limited [nutrient] factor.” es (8 to 16 inch depth). Results Commonly illustrated with a wa- didn’t show significant difference ter barrel and staves analogy, the between the two horizons in our maximum water volume (repre- clay loam soil and my frugal side senting cumulative plant growth) couldn’t justify doubling the cost. cannot exceed the shortest stave Although I subsoil and deep till (limiting nutrient) regardless of the ground before planting I’m the height (abundance) of any assuming the affect on nutrient other stave (nutrient). Implied is mixing is minimal. Admittedly, that rectifying this nutrient short- it would be better to sample af- age will advance growth but only ter the finish tilling especially in up to the level of the next limit- terms of knowing the levels of the ing nutrient. Further, inputs of soil’s less mobile nutrients like any non-limiting nutrients won’t phosphorus, potash and lime. improve yield but pointlessly Constituting 100% of our labor increase costs of production. force, I don’t always get every- Herein lies the value of soil test- thing done on time or in proper ing. Finding a testing lab is easi- order. ly done with an online search or request to the state university ex- My practice is to scalp each sam- tension service. Two items need ple site about a ½ inch down to consideration: sampling meth- mineral soil, removing identifi- od and interpretation of results. able plant debris, then using a Both are well addressed online. soil probe to remove an eight inch core. About 20 cores are ran- Sampling for soil analyses pres- domly collected into a stainless ents the typical sampling chal- steel dishpan, (Don’t tell Mom!), lenges. How many samples are then left covered to dry, sifted necessary and how/where shall through a ¼ inch or smaller sieve,

398-8 thoroughly mixed, and a lab de- I always request the lab’s option fined volume sent for analysis. of graphically displayed results My homemade probe was made in addition to the regular tabular from a couple feet of thin-wall data. These bar charts plotted chrome plated tubing from a against recommended ranges mop handle. (Again, don’t tell you are very useful in visualizing the know who!) A short length of PVC degree of excess or deficiency for pipe with a tee handle drives the each nutrient. Once again, self tubing into the soil, and a longer education online is essential in length of PVC is used to eject the understanding the test results cores into the dishpan. Absent a and planning remediation. probe, samples can be taken with a clean (rust free) spade or shovel While lab tests drive our initial by removing a soil slice, then cut- field management, much of our ting an 8” vertical column from ongoing practice is based on ex- the slice. Care should be taken to perience and common sense. create uniform columns from site What follows is a “seat of the to site. pants” philosophy that works well for us.

• Nature is benevolent. Even without intervention plants grow, soils produce, fertility exists. Plants, especially peonies, have evolved superb abilities to scavenge nutrients within their rhizosphere. For us it is not entirely necessary to achieve 100% growth potential. Do what works, don’t get greedy. • Recognize the TWO soil types. Soil mixes in potted stock are crit- ically different than the soils of field grown plants. With all of their roots in a pot, there is no escape for mistakes made. Field grown stock is more forgiving of poor decisions. (All comments in this article are applicable to field growing. Watch for an upcoming article on potting soils.) • Many off the shelf (home gardener) proprietary fertilizers recom- mend rates that approach near toxic levels. Less is more! When lacking a lab-based soil test, use the Rule of Half. You can always apply a second ½ dose later. • Know some Chemistry. It’s FUN! Begin by memorizing about 35 of the most familiar atomic symbols from the first four rows (Pe- riods) on the Periodic Table especially those in the first and sec-

398-9 ond columns (Groups/Families) and within the four same periods for Groups 13-17. Ignore all of groups 3-12 and group 18. This is the lexicon for thinking chemically about nutrients. Don’t mem- orize any of the data. (It’s on the #*%$ table and you won’t need it anyway!) Next, learn the essence of ionic bonded salts (i.e. fertilizers), positively charged cations (from groups 1 or 2) held to negatively charged anions (from groups 16 or 17). This will help make sense of the Cation Exchange Capacity (CEC) on soil tests. Likely you’re familiar with most of this already. • Learn to recognize visual (foliage) symptoms from online photo libraries.

Keep in mind the following nu- N sources. More recently, urea, trient features when adjusting CO(NH2)2 (46-0-0), has nearly re- fertility: placed ammonium nitrate. Both (and other ammonia/ammoni- Nitrogen: N is typically the first um types) tend to drive down soil and most limiting nutrient in pH with repeated use. Nitrogen temperate soils due to it easily strongly promotes vegetative leaching beyond the root zone. growth and plant juvenility. Care- Irrigation and precipitation ex- less or excessive applications of acerbate this. Lacking a posi- N can ‘burn’ plant tissues by salt tive charge in most forms, it is desiccation. Ammonium forms not attracted to soil particles. within the plant need sunlight to Some forms such as ammonium break down. Low levels of light NH4+ and nitrate NO3- (the forms in greenhouses, especially in plants can immediately use) do the short days of spring, may re- have electrical charges. Soil or- sult in toxic accumulations from ganisms use (sequester) N when this form. Growers prefer nitrate they feed on carbon. Higher forms at this time of year. levels of organic matter in soils helps retain N in the bodies of Phosphorus: P is in many ways microfauna. Farm them by split- the antithesis of N. It drives plants ting N applications as soil warms towards maturity and reproduc- in the Spring. tive phases for flowering, seed, fruit and root production and is Historically, ammonium nitrate, valuable to work into soils before NH4NO3 (34-0-0) was a prevailing planting. It tends to be tightly commercial N source and was held to soil particles, especially useful in that both halves are in clay soils with high cation ex-

398-10 change capacity, CEC. As a result K at planting time. Procuring K P leaching is minor from precipi- in other than triple NPK formu- tation and irrigation. On the oth- lations (10-10-10 or 12-12-12 or er hand, top-dressing P is less ef- 16-16-16) is a challenge for us. fective than incorporating it into Locally, “K-Mag” (aka SulPoMag the root zone because it tends to when I knew it in Michigan) is stay where you put it. nearly the exclusive form available here in orchard country. Compared to N it is very safe Sulfate of Potash Magnesia is regarding toxicity to the plant. 0-0-21+21S+10 Mg, and is only Bonemeal, and rock phosphate partially soluble in water. It does are moderate sources of phos- not affect soil pH. Potassium Ni- phorus and are often used as trate, KNO3 (13-0-44) is entirely bulb fertilizers. Treble super- utilized for both its K+ cation and phosphate (0-45-0), is a high NO3- anion. Completely soluble, phosphorus form, and our favor- useable for foliar application, it ite for fall incorporation at plant- is expensive and mostly used as ing time due to its high P and a glasshouse fertilizer. lack of N which would mostly be leached out over winter. We also Historically a major component use monoammonium phosphate of colonial gunpowder, it became or “MAP” (11-52-0), in spring, as nearly impossible to source after its easier to obtain in our area 9/11. Most economical and avail- than superphosphate and a able is Muriate of Potash “MOP” moderate source of N. There is Potassium Chloride, KCl (0-0-60), also double ammonium phos- the most widely used K fertilizer. phate “DAP” but less commonly Its chlorine may be detrimental available (for us) than MAP. especially in soils already high in chlorine or in areas with high Potassium: K (from the neo-latin chlorine irrigation water. Typ- “Kalium”), is the underappreciat- ically it is generally not a con- ed stage manager, excluded from cern except for chlorine sensitive the curtain call. K is not a major crops. Another K source is Sulfate component of plant tissues but of Potash, ”SOP” K2SO4 0-0-50, is the catalyst for a multitude of which provides sulfur in addi- physiological actions and chem- tion to its K and is good for chlo- ical reactions. It is a Group 1 cat- rine-sensitive crops, but is more ion, well held or exchanged by expensive. Importantly, the “Salt high CEC soils. Although more Index” for MOP is 116 but for SOP soil mobile than phosphorous, it only 46. Nonetheless, the salinity is beneficial to also incorporate

398-11 risk is low for almost all crops un- your soil will require to provide a der normal irrigation. long term effect. Swatting a mos- quito solves an immediate prob- pH: Acidity vs. Alkalinity. Nu- lem. Knowing how many more trients are most available when are coming out of the woods en- soil pH is between roughly 5.5 to ables long term strategy. 7.5 moderately acidic to nearly neutral/slightly alkaline. Garden Quick Lime, Calcium Oxide, vegetables prefer a higher near CaO: This is the brilliant white neutral 7.0 pH soil while many lime powder used cautiously in landscape species trend towards masonry to make mortar. It is moderately to slightly acidic, very caustic (the opposite of “cor- 5.5 to 6.5. Lactiflora type Paeo- rosive”). “Quick” is the colloquial nia do well in this range but will term for alive, as in “the quick maximize growth in the nearly and the dead” or the “quick” of neutral 6.5 range. The Ericaceae a fingernail. It’s hard on tools, family, Heaths, Heathers, Rhodo- sinuses, skin and plants. Adding dendrons etc., are famous acid water must be done carefully as lovers having evolved to thrive at the exothermic reaction releas- the lower pH of peat soils. Each es intense heat. The pH raising whole pH number is an order of power of liming materials is due magnitude (10 X) greater or less- to the anion, in this case the ox- er than the next whole number. ygen in Quick Lime. (CCE about The most significant effect of 175%.) extreme pH is the reduced avail- ability of soil held nutrients and Slaked Lime, Calcium Hydrox- the increased release of toxic el- ide, Ca(OH)2: Also quite bright ements. looking, this form is less caustic or dangerous to handle than Lime and Liming: This term is Quick Lime. Slaked, as in slaking about as specific as “sandwich”. one’s thirst, refers to the addition The main role of liming is to raise of water to Quick Lime. This hy- the pH of acidic soil thereby mak- drates the lime converting CaO ing nutrients more available. A to Ca(OH)2. Again, it’s the anions, secondary goal can be to supply two hydroxide OH’s that provides extra calcium, Ca (or sometimes the pH increase. (CCE about magnesium Mg as well). Several 135%.) distinct chemicals are termed “lime.” The “pH buffer capacity” Ground Limestone: primarily on a soil test is essential to un- Calcium Carbonate: CaCO3 is ‘Duchesse de Nemours’ after frost; not much cansourced be done about from the sedimentaryunsightly unsightly oce leaf - derstandingdistortion. Disease the may degree set in. of liming

398-12 anic rock and can vary in col- easier to find than ground lime- or from slightly tan to gray. It is stone. (CCE about 75-105%.) very mild compared to Quick or Slaked Limes. CCE is theoretical- Lime Score: The pH raising abili- ly 100%, but varies from 75-95%. ties of limes are quantified based The most commonly sold garden on calcium carbonate equiva- lime, it is often misrepresented lent, CCE. Pure calcite mineral as Dolomite or Dolomitic Lime- CaCO3 is set as 100%. Several oth- stone (and vice versa). er systems exist which include the effect of moisture content Dolomite strictly refers to the and particle size of the material. mineral Calcium Magnesium Fine particles raise pH faster, but Carbonate: CaMg(CO3)2 and con- persist less than coarse particles. tains two anions of carbonate Lime, like phosphorus, is much plus magnesium Mg as well as more effective when incorpo- calcium. Seldom available, but rated into the top 6 to 8 inches often misspoken for Dolomitic of soil. Again, excellent resourc- Limestone. es are available through online state extension websites. Dolomitic Limestone is a vari- able mixture of both CaCO3 and This just scratches the surface, CaMg(CO3)2. The packaging gives but I hope is a useful beginning. the percentage of each. It is often

To find soil testing services by state, visit: gardeningproductsreview.com/state-by-state-list-soil-testing-labs- cooperative-extension-offices/

398-13 If Wishes Were Peonies The APS’ 2021 Virtual+ Convention

Throughout the past year, the wish of the American Peony Society has been to hold our 2021 convention live in Minnesota. So we are particularly saddened to cancel this year’s event due to COVID -19 pandemic precautions, especially knowing so many members have worked hours planning the convention and preparing tour gardens. Thanks go to David and Linette Sorrentino, and the Minnesota Peony So- ciety, for all of their efforts to provide Membership with an outstanding convention experience. APS is hopeful their plans may be put to use for a future event in Minnesota! Plan B’s a Little “Un-Conventional” Knowing our members need a “PEONY FIX” more than ever, the APS has put together an online convention line-up of tours, seminars, meetings and a virtual peony show! This year’s virtual convention is titled “IF WISHES WERE PEONIES”. If wishes were peonies, we’d probably all need to buy more property, but the online options should help to alleviate this possibility! We’re learning new ways to host a convention, revealing many silver lin- ings to the initial cloud. The virtual format allows us to extend activities through spring and summer, and make the experience available to members around the world. Our virtual APS Convention now is more open, more accessible, and more affordable than ever (most activities are free). There’s also a regional “live” twist: we’ve connected with our commercial and other members to highlight live in-person tours and garden events. Find Details & Dates on the APS Website Convention activities begin in mid-May and proceed through mid-Au- gust. Public options may be found on the APS website’s public pages at: https://americanpeonysociety.org/news-events/if-wishes-were-peonies/ or under the News and Events tab. Member only options (you will need login) can be accessed on the Member Portal by clicking on If Wishes… in the Portal’s main navigation and selecting a detail tab. Signup and submission for Member offerings can only be made through the Member Portal.

398-14 Three Months of Amazing Peony Experiences 1) The Photo Contest & Virtual Show. Join us for a virtual photo contest/flower show, which allows any Member to make submissions of peony related images to the website for display in an online gallery, showcasing a number of classes. Share your love of peonies through submitting images for the world to see and enjoy “light” competition with fellow Members. Winning entries will be featured in the APS Bul- letin, social media and other outlets. 2) Virtual Seminars. Our popular seminars will now be offered through Zoom and will surely be of interest to those who enjoy hybridizing, historic peonies of Minnesota, cut flower production or simply want to know the basics! APS has elected to make these “member only” as the format is new to us. 3) Live Tours. Tours of gardens and commercial growers’ operations are available for the general public and are listed on the public side of the APS website. Additionally a number of Member gardens in the Minne- sota-Wisconsin area will be open for APS members only. These member gardens require signup through the Member Portal. Join in the beauty of live peonies if you are able. These Member gardens would have been on tour for the cancelled convention and the owners are looking forward to showing off their work. 4) Video Tours. Share your gardens with a video tour! APS would love to make your gardens available through the Member Portal: submit your video throughout the spring and summer months. 5) The Annual General Membership Meeting. Our annual meeting will be held via Zoom, and include the “State of Union”, Personal and Gold Medal Peony Award announcements, election conclusion and basic Society Information. 6) SPONSORSHIP! If you are able to contribute to help defray costs with “If Wishes Were Peonies” APS would appreciate your generosity. Sponsorship has a number of giving level and APS will gladly list you as a donor. Join us! You’ll find all the Convention activities and information at americanpeonysociety.org/news-events/if-wishes-were-peonies/

398-15 Paeonia ssp. veitchii 'alba'

Paeonia arientina 'dark'

Paeonia banatica

Paeonia intermedia

Paeonia mairei 398-16 Paeonia peregrina

Paeonia ostii

Paeonia arientina 'dark'

Paeonia potaninii v. trolliodes Paeonia potaninii

Paeonia qiui

Paeonia tenuifolia 397-17 The Language of Peonies

Carpel [kär-p l]e

Originating from the Greek word Karpos, meaning “fruit”, the word carpel is used in peonies to describe the female flower structure enclosing the ovules which may later become seed. Carpels, fin- ger-like structures, grow in size as they mature and may number from one to many depending upon genetic origins of the plant.

397-18 Th e 2021 Photo Contest & Virtual Exhibition Share, Display and Enjoy Your Blooms in a Stunning Virtual Gallery

Cate Crawford - Exhibition Co-Chair Th ough our Annual APS’ convention is online this year, APS isn’t letting that stop us from celebrating the exquisite beauty of the peony! Th e APS is thrilled to announce the fi rst-ever APS Photo Contest & Vir- tual Exhibition. As part of the 2021 APS convention - “If Wishes Were Peonies” - the Vir- tual Exhibition will provide all APS members a chance to showcase our beautiful gardens and highlight the special love we share for our peonies. As part of the exhibition, we invite all APS members to submit peony and garden photos in a broad range of categories including but not limited to: Cultivars Arrangements and Bouquets Full Plant Many More! APS will display these member photos on their website for all to appreciate. APS is also assembling a Member Judging Panel who will pick their favorites and assign awards! Photo submission kicks off May 15 and extends through August 1. Sub- missions can be uploaded via the APS Member Portal. Judging will run Aug 15 - Aug 31, with winners, awards and Court of Honor announcements to follow on Sept 10. Photos of particular merit will also be showcased in APS publications and events going forward. All APS members will receive a kick off email just prior to the Virtual Exhibit opening. Th e kick off will make sure members 'Magical Mystery Tour' have all the logistical informa-

398-19 tion required to submit photos. APS will also announce the start of the competition via social media. We are still assembling the Virtual Exhibition Judging Teams. So, if you’re interested in participating please shoot an email to Cate Crawford at [email protected]. Given the gardening prowess we know exists within the APS membership, we look forward to a bounty of stunning images we can all enjoy. In a time de ned by separation and struggle, our sincerest hope is that the Virtual Exhibition provides a beautiful, immersive experience - connecting us to peonies and to each other.

Photos, Nick Maycher

'Rose Heart'

'White Innocence'

Learn more at: americanpeonysociety.org/news-events/if-wishes-were-peonies/ #virtual-show Submit Photos via the APS Member Portal at: americanpeonysociety.org

398-20 Leena Liljestrand, APS Director, President of the Swedish Peony Society - Mariestad, Sweden

What happens, when a peony and showed them her plans to enthusiast's garden gets far too create a peony park of this for- full of all kinds of shrubs, trees, gotten area in the middle of this and perennials, not to mention small town. After an all-positive hundreds of peonies? She enjoys meeting, the only thing left was it all wholeheartedly for a while, to roll up the sleeves and start! but then her eyes start to wan- der to the other side of the fence, This is the story of The Village where a 1.5-acre communal Park in the small municipality wasteland, covered with couch of Mynämäki in southern Fin- grass, screams for TLC. This was land, half an hours’ drive from in 2017, and in 2018 she went to the city of Turku. In this park the meet the decision-makers in the peonies are the stars, and the town hall, and both described enthusiasts giving them perfect

398-21 surroundings to shine are Sari kind-hearted and effective Kau- Kantelo, a board member of The ko Kylänpää have been at work, Finnish Peonists, and her hus- making it possible to start build- band Juha Laine. The goal is to ing the flowerbeds and planting create an experience for all visi- the trees, which will give some tors, whether they are town res- well-needed shade when the hot idents, hobby gardeners, tour- summer sun is shining. ists, or garden professionals; to show everyone the diversity of Everything is done with sus- peonies, how they grow in differ- tainable development in mind, ent locations and soils, and how using recycled and eco-friendly they match and thrive with oth- material, not only because of er perennials, trees, and shrubs, the planet but also the wallet, creating pleasant entireties. as Sari and Juha pay the costs from their own pockets. The pe- The whole project was so onies are divisions of Sari’s own mind-blowing that it took some plants, as well as donations from time for everyone around Sari to friends and fellow members. understand that she was quite Young trees have been donated serious. The residents around by a local nursery, the Munici- the area were first confused, pality of Mynämäki has donated then overjoyed realizing how water posts and material for the much beauty would be growing many paths in the garden. And in the unexploited area. People of course, The Finnish Peonists passing by have already noticed support the project in various that there are so many more but- ways. terflies and bees around, thanks to the park. Some have followed Today more than 300 different the activities from their apart- peonies have been planted, and ment buildings, saying that they when everything is finished, the broke in sweat just watching the number will be around a thou- amount of work the enthusiasts sand. It is not the number that is were doing daily. They added important, but the quality, says that it is like watching a mix of an Sari. There are to be species, action movie and a documenta- hybrids, lactifloras and inter- ry – the garden being built plant sectional peonies, maybe some by plant, stone by stone, and hardy wooden ones as well. The that it is very inspiring! fenced-in park will have specific opening hours and will be not The work started three years ago. only a feast for the eyes but also The excavator and its owner, the a place for studies, contempla-

398-22 398-23 tion, to enjoy a moment of peace going project in Finland, in the and harmony amidst all asphalt. city of Kotka, about a 1½ hours' Lectures about peonies – free drive to the east from Helsinki, from charge – are planned when in a city-owned and maintained everything is ready, and we are park. This fall approximately quite certain that this lovely, ver- 200 peonies were planted, 15 of satile Village Park will be a source each cultivar. The plan is to plant of inspiration for every visitor around a thousand peonies in and will increase the knowledge the City Park of Kotka in coop- of the different peony species eration with The Finnish Peony and cultivars, how to plant and Society. The peonies are donat- care for them. As Sari and Juha ed by the members and a peony say: Shared joy is a double joy. nursery in Holland. The mem- We will follow the development bers are to take care of the main- and tell you more about this fan- tenance of the peony park. More tastic project in the future. about this you can see here: https://www.facebook.com/pe More photos can be seen on Ins- onyparkkotkafinland/ tagram: https://www.instagram. com/pionipuisto/ Photos: Sari Kantelo, Board Member, the Finnish Peonists More peony news from Finland: There is also a newer (2020), on-

398-24 PART ONE

Donald R. Smith – Fairfield, Connecticut

The history of Pæonia is a very George Anderson presented to long one, with some recent es- the Linnaean Society of Lon- timates placing the origin of the don in 1818, which described 13 genus in the Cretaceous period species of Pæonia. This was fol- of the Mesozoic era between 80 lowed by a more comprehensive and 110 million years ago (mya). study published in 1890 by Rich- Until quite recently, little was ard Lynch, which recognized known about the evolutionary 25 species and was the first to history of the genus. However, divide the genus into 3 subge- recent DNA sequence studies of nera. This classification stood Pæonia have provided the first until 1946 when Frederick Stern significant information from published his seminal study on which reasonable estimates of the genus, “A Study of the Ge- some of the major evolutionary nus Pæonia”, which described events of the genus could be 33 separate species. Stern’s tax- made. One example is the ap- onomy system was long consid- proximate timing of the first ma- ered the definitive work on the jor evolutionary divide, the split genus Pæonia and as such has of section Onaepia from the re- stood relatively unchallenged mainder of the genus, which is for more than 50 years. estimated to have occurred in the middle of the Miocene ep- This situation began to change och of the Tertiary period during in 1997 with the publication of the Cenozoic era somewhere the first of a series of landmark between 15 and 20 mya. papers by Tao Sang et al. re- porting on the results of DNA se- The first written records of the quence analysis of 32 species of genus date back to the first Pæonia. These studies revealed century A.D., but the first thor- numerous surprising discover- ough study devoted entirely to ies that laid the foundation for Pæonia was a monograph by a greatly improved re-classifica-

398-25 tion of the genus. The two most considerable room for differ- significant findings of these ences of opinion and frequent seminal studies are summa- debates among taxonomical rized below. experts. Classification has been particularly difficult in peonies Perhaps the most important and other plant groups which finding of this series of stud- have undergone extensive re- ies was that the majority of the ticulate evolution. These diffi- species in section Pæonia were culties have resulted in numer- determined to be of hybrid ori- ous adjustments, changes, and gin, a conclusion that has major reclassifications over the years. implications for the classifica- Recent advances in modern ge- tion of the genus. However, the netics, however, have made tax- biggest surprise to come from onomy a much more precise sci- these studies was the unexpect- ence with significantly less room ed discovery that many of these for error and ambiguities. For hybrid species had speciated example, the recent application without elevating to the tetra- of molecular markers has great- ploid level as is usually the case ly facilitated the detection of hy- in most other genera. bridization and the recognition of allopolyploids (having two or Creation of new species (specia- more sets of chromosomes from tion) through natural hybridiza- different species) in many plant tion, especially when combined groups. This is especially true in with polyploidization, is an im- Pæonia, where the recent land- portant mechanism in plants mark studies of Dr. Tao Sang and it has made the science of and his colleagues have clear- classifying plants (taxonomy) a ly established that reticulate very challenging field of study. evolution (hybridization) has This mechanism is known as played a primary role in specia- reticulate evolution as it is best tion in peonies. Starting in 1995 described as a network rather with his doctoral dissertation at than as a branching tree. Ohio State University entitled Until quite recently, nearly all “Phylogeny and Biogeography plant classifications were based of Pæonia (Pæoniaceae)”, Dr. solely on careful examination Sang has laid the foundation of morphological characteris- for a completely new and much tics and differences. As a re- more accurate reclassification sult, taxonomy has remained a of genus Pæonia, and especial- relatively inexact science, with ly section Pæonia (i.e., Pæon),

398-26 where natural hybridization paternal parent by the process has been particularly prevalent. of elimination). Although the details of the science and methods used in these By analyzing both nuclear and studies are extremely complex chloroplast DNA sequences, and technical, the results and Sang and his colleagues were conclusions can be more easily able to accurately reconstruct summarized and described in the complex reticulate evolution laymen’s terms. Thus, I will at- patterns within section Pæonia. tempt to do this here, in the next However, they also emphasized few paragraphs. that this reconstruction may still be an underestimate of the The analysis of DNA sequences natural hybridization which has can be an extremely powerful occurred in this group. Yet, as tool in reconstructing reticu- mentioned earlier, the most sur- late evolution. Sang analyzed prising conclusion from these two different types of DNA se- remarkable studies was, un- quence data to help reconstruct doubtedly, the high proportion the very complex pattern of of diploids seen among the hy- reticulate evolution in section brid species. Although the tet- Pæonia. First, he analyzed nu- raploid species have often been cleotide additivity detected by suggested to be allotetraploids directly sequencing PCR prod- (thus indicating hybrid origin), ucts of internal transcribed none of the diploid species of spacers (ITS) of nuclear ribo- this section had ever previously somal DNA. This method can be been considered to be of hybrid highly informative and accurate origin. in detecting hybridization in cases where parental sequenc- A summary of the major capabil- es are maintained in the hybrid ities of this technique are given species, which is the case in below. They can: peonies due to the slowing of concerted evolution caused by 1. Detect hybridization events the long generation time of pe- (including relatively ancient onies. Next, Sang analyzed the events and multiple hybrid- rapidly evolving (maternally in- ization events in the same lin- herited) chloroplast gene, matK. eage). This technique can identify the 2. Identify the parent species of maternal parent of a hybrid spe- each hybrid species. cies (and thus when combined with ITS sequence data, also the

398-27 3. Identify the maternal and lia, P. veitchii and P. intermedia, paternal parents (including are not derived from hybridiza- extinct species and ancient tion. ancestors) of many hybrid • The existence of three extinct species. ancient species in section 4. Identify closely related spe- Pæonia was determined from cies that share a common par- their DNA sequences observed entage, thus forming a sister in their extant hybrid descen- group. dants, which have supplanted 5. Determine the relative order them. in which various hybridization • Extensive reticulate evolution events occurred. in section Pæonia has made 6. Determine the approximate classification of this section time when a certain hybrid- extremely difficult. ization event took place (i.e., • Most hybrid species are found determine the time of origin in the Mediterranean region, of the new species). whereas their parental species are presently restricted to Asia. By applying these state-of-the- • The earliest evolutionary split art techniques to the study of in genus Pæonia probably oc- the Genus Pæonia, Sang and his curred between section Onae- colleagues were able to make a pia and the rest of the genus, number of significant discover- thus making Onaepia ies concerning the evolution of the old- est section. It is estimated Pæonia. A summary of the most that this split occurred approx- important conclusions from imately 15 million years ago. these studies are listed below.

• The majority of the species in The results of Sang’s phyloge- Pæo- section Pæonia are of hybrid netic analysis of section nia origin. are summarized in Figure 1, (which is an updated and rear- • The proportion of diploids ranged version of figure 7 from among the hybrid species is the initial published paper by surprisingly high, suggesting Sang et al.1 in 1997). By exam- that hybrid speciation at the ining this figure, it is possible diploid level has been quite to trace the reticulate evolution successful in peonies. within section Pæonia by follow- • Only four species in section ing the evolutionary tree from Pæonia: P. lactiflora, P. tenuifo- left to right.

398-28 Figure 1. An updated phylogeny chart for section Pæonia, based on the DNA sequence studies of Sang et al.1,2,3

In ancient times, following the also between members with- initial split of section Onaepia in the large clade. Many of the from the rest of the genus and resulting hybrids survived and the subsequent split of sections evolved into new species, which Moutan and Pæonia (which are in a few cases replaced one of not shown in the figure), section their parental species. Some Pæonia evolved into two distinct accomplished this through poly- clades, a large clade (top of fig- ploidization, while others man- ure) and a small clade (bottom). aged to survive by remaining at The original ancestors of these the diploid level. This proba- two clades hybridized to create bly occurred through extensive a new hybrid species, which in vegetative reproduction by rhi- turn, replaced its paternal par- zomes which facilitated survival ent to become the only surviv- of the initial diploid populations ing ancestor of the small clade. of hybrids until they became fer- After this, extensive hybridiza- tile or underwent polyploidization occurred between various tion. members of the two clades and

398-29 Figure 2a. The initial classification of the Genus Pæonia based solely on the DNA sequence studies of Sang et al. 123. After careful review of the vari- above (see Figure 2a). This ma- ous papers mentioned above, I terial was first described in a was able to construct a prelim- number of articles in the Pæo- inary new classification of Pæo- nia newsletter (Pæonia, V30N3, nia which incorporated all of the V30N4 and V31N1), beginning in essential conclusions and results the summer of 2000. of the DNA studies described

398-30 Figure 2b. The classification from the recent book “Peonies of the World, Taxonomy and Phytogeography” by D-Y Hong 4. In 2004, the results of a new the genus. At the time, this book comprehensive taxonomical was thought of by many (in- study on peonies by Josef Hal- cluding myself) as a possible re- da were published in the book, placement for the famous Stern The Genus Pæonia, which also monograph discussed earlier. I contained a new classification of reviewed this book in an article

398-31 in the December 2004 issue of ysis of his own morphological the APS Bulletin (No. 332, p.224- data. Although this new classifi- 227), in which I compared Hal- cation appeared to be a clear im- da’s new classification with that provement over the earlier Stern of Stern and also with my prelim- classification, there were several inary classification based on the differences and inconsistencies DNA studies mentioned above. with my classification based pri- Based on this comparison, I was marily on the DNA results of Sang compelled to reject this new et al. Thus, we were left with the classification by Halda, because dilemma of having two new clas- it was in serious disagreement sifications of Pæonia based on with both the DNA results as well two comprehensive and equally as with the long-standing classi- credible studies using two com- fication of Stern. In May of 2005, pletely different scientific tech- I presented these results and niques that were not in complete comparisons at the Internation- agreement. And, unfortunately, al Peony Symposium in Munich, some of these differences and Germany. Then, in the spring of inconsistencies were fairly sig- 2010, came the long-anticipat- nificant and, therefore, could not ed book by De-Yuan Hong sum- simply be ignored. marizing the results of a 25-year study of the genus, that was the Thus, I began a new study spe- most comprehensive and thor- cifically directed towards identi- ough study ever conducted of fying and resolving these incon- the genus Pæonia. After a quick sistencies. My initial approach review of this book, it appeared was to give a much higher cred- to be the long-awaited replace- ibility to the DNA results, since ment for the Stern monograph. DNA analysis is known to be a very “exact” science compared However, a more extensive and to the more subjective science of detailed review of this book re- taxonomy. Therefore, whenever vealed that, although Hong was the two results were in obvious aware of the DNA studies of Sang conflict, I would simply choose et al., he did not use or incorpo- the more precise (DNA) result rate any of the findings into his over the inherently less accurate analysis and classification. Un- one. It took me several years to fortunately, no explanation or get off this track and to finally re- reason was given for this crucial alize that both techniques have decision. As a result, his pub- their separate advantages and lished classification (see Figure also, more importantly, share a 2b) was based solely on the anal- few common potential sources

398-32 of error that are completely in- beled. His argument and sup- dependent of the inherent accu- porting evidence seemed com- racy of the technique or method pelling, so I quickly looked into employed. It is this latter point this matter and came away con- which eventually brought me to vinced he was probably right. a satisfactory resolution of this His claim was that the sample problem. identified in the Sang study as (P. russi) P. russoi was probably Although DNA sequence analy- mis-labeled and was instead ac- sis is an extremely precise sci- tually P. corsica, since P. russoi is ence, problems can arise from not known to grow in the region mis-identified or mis-labeled of the western Mediterranean is- samples, just as in any field of lands, listed as the collection site science. For this reason, it is very of the study sample. The only important that the precise loca- peonies known to inhabit these tion of the collection site for each islands (Sardinia and Corsica) sample is accurately known and are a species known as P. corsi- recorded. Taxonomic studies are ca, which is closely related to P. needed to identify the species to cambessedesii. P. russoi, on the be studied and where to collect other hand, is only found in Sicily them. Even then, the geograph- and the southern part of Italy. P. ical distributions of species russoi is considered a subspecies sometimes overlap each other. of P. mascula to which it is close- Therefore, the morphological ly related. In addition, it was characteristics of each sample quite apparent that making this need to be carefully checked change would also eliminate one to assure proper identification. of the more troubling discrepan- Many closely related species are cies between the two classifica- quite similar in appearance, and tions. This knowledge sent me in thus, can easily be mixed-up or a new direction in search of pos- mis-identified. sible solutions to other disagree- ments in these classifications. This problem was first brought to my attention by an email I re- Sometime after the publication ceived from John Hudson, (the of Hong’s book, I became aware editor of the Newsletter of the of a paper by Pan, Zhang and Peony Group of the Hardy Plants Sang7 published in 2007 in the Society, UK), informing me he American Journal of Botany, was convinced that one of the presenting evidence for the or- samples measured in the Sang et igin of a diploid hybrid species al. studies was probably mis-la- in Pæonia. In a note under the

398-33 section “plant materials” the au- have been a mix-up in the iden- thors pointed out several recent tification of any of the samples nomenclature changes affecting used in the DNA sequence study, the four Pæonia species investi- but especially with the one iden- gated in their study. One of these tified as P. anomala. changes was that P. sinjiangensis (which is sometimes spelled as Unfortunately, there were no P. xinjiangensis) had been re- precise coordinates provided named P. anomala. It was clear for the samples studied. The after reading this article that the only type of information given main species (P. anomala) they on the location of the collec- were studying was the same spe- tion site was the county and the cies, which Sang et al. had stud- province. Although this type of ied previously under the name information is generally suffi- P. xinjiangensis. Except now, it cient to unambiguously identify was being called P. anomala. It most species, in this case it was seemed very simple to correct not. Nonetheless, it quickly be- this problem by simply chang- came apparent that there are ing the label on my charts to two fairly similar species which reflect this change in nomencla- have distribution ranges that ture. However, doing so would partially overlap, where a col- create a much larger problem, lected sample might easily be since there was already anoth- mis-identified. These two spe- er sample included in the study cies are P. anomala and P. inter- that was labeled P. anomala. media, both of which are native Moreover, since these two sam- to and widely distributed in the ples gave very different DNA se- Altai area of Xinjiang province in quence results, they were clearly northwestern China. Since the not the same species. Thus, if exact locations of the collection P. xinjiangensis was actually P. sites of the samples were not anomala, then the sample orig- provided in the published paper, inally labeled P. anomala had I used the approximate locations to be something else. This led given in order to assess the like- me back to the original paper lihood of an identification prob- by Sang et al.1 to check for the lem. It soon became clear from collection site locations for the a careful examination of these measured samples and then to distribution maps that both of the geographical distribution the samples in question (P. xin- maps of the various species in jiangensis and P. anomala) were Hong’s book to see if there could collected from sites that were located in or near the edge of an

398-34 overlap region in the distribu- P. intermedia, which was simply tion ranges of P. anomala and P. mis-labeled as P. anomala. intermedia. The collection site for P. xinjiangensis (XIN) was in Once these two important the middle of a large area of the changes/corrections were made known distribution range of P. to the initial version of the phy- anomala, but also near the edge logeny chart derived from the of the known distribution range DNA studies, all of the most trou- of intermedia. The collection bling discrepancies between site for P. anomala (ANO) was these two different methods in the middle of a known dis- were eliminated, leaving only a tribution area for P. intermedia, few relatively minor issues to be but also close to the edge of the resolved. known distribution area for P. anomala. Therefore, the identity These remaining issues are sum- of these two samples could not marized and discussed below: be unambiguously determined 1) Is P. veitchii an original ancient based solely on their collection (non-hybrid) species (Sang) site locations, even if these sites or a sub-species of P. anomala were precisely known. Howev- (Hong)? er, in either case, there are only two possible choices, P. anom- 2) What is the proper placement ala or P. intermedia, since there of P. sterniana and P. banati- are no other species of peonies ca? known to populate these two 3) What is the proper species sta- regions. However, since the DNA tus and placement of P. humi- sequence signatures for these lis? two samples were distinctive- 4) Are P. mlokosewitschii, P. rus- ly different, they cannot be the soi, P. wittmanania, P. banat- same species. Therefore, one of ica, etc. separate species these samples must have been P. (Sang) or subspecies (Hong)? anomala and the other was then P. intermedia. The paper by Pan 5) How many sub-sections are et al. clearly established that P. necessary to classify section anomala is the hybrid species Pæonia? 2 or 3? that resulted from a natural cross between P. veitchii and P. Although there were a number of lactiflora, thereby also establish- different considerations which ing that the other sample was factored into the final resolution of these remaining discrepancies, there was a single overrid-

398-35 ing principal which was invoked. phological similarity to P. veitchii Since I have no further evidence (see Figure 1 and Table 1). or suspicions that any other samples may have been incorrectly 2a) There is a good case to be identified or labeled, I chose to made for placing P. sterniana use DNA results, whenever they with P. emodi (which is its ma- were available, over morpholog- ternal parent) as Hong has done. ical data. On the other hand, the However, I believe that there is a morphological data were invalu- more compelling case for plac- able for determining the proper ing P. sterniana with its paternal placement of all of the newly dis- parent, P. mairei, and alongside covered species or other species P. banatica which also shares the or subspecies not included in same paternal relationship with the DNA study. With this general mairei. guideline in mind, the resolution of each of these five remaining 2b) Likewise, although the DNA issues is discussed below. data indicates that P. banatica and P. officinalis share one par- 1) The DNA data indicates that ent (Ari-Par Group), P. banat- P. veitchii is one of only four ica also shares a parent with non-hybrid species in section P. sterniana and the Cam-Cor Pæonia and that P. anomala is sister group and thus a strong a hybrid species from a natural argument can be made for plac- cross with P. veitchii being one ing P. banatica with these other of the parent species. There- species and its paternal parent, fore, there is no DNA support for P. mairei, because they are all Hong’s conclusion that P. veitchii interclade hybrids as shown in should be considered as a sub- Table 1. species of P. anomala. In fact, P. humilis it would seem more appropri- 3) Hong lists as a sub- P. officinalis ate and justifiable to consider species of . Howev- P. anomala as a subspecies of P. er, based on the DNA analysis P. humilis veitchii rather than the other way of Sang et al., has the P. arietina around. This is especially true in same parentage as P. parnassica P. of- light of the suggestion by Pan et and (and not ficinalis al. that P. anomala may actually ) and therefore, should be a second-generation hybrid be grouped together with these species (resulting from a further other two species as the third back-cross with P. veitchii), thus member of the Ari-Par sister P. officinalis explaining its high level of mor- group. , on the other hand, was determined to be the

398-36 progeny of a natural cross be- ent grandparents including two tween P. peregrina and a mem- that are now extinct. Although ber of the Ari-Par sister group, P. daurica might well be one of which includes P. humilis. This its parents, P. wittmanniana’s DNA data also suggest that P. hu- unique parentage places it mid- milis might be both a grandpar- way between P. obovata and ent and a parent of P. officinalis the clusii – mascula sister group which supports Hong’s conclu- (which probably includes P. dau- sion that these two species are rica) and clearly justifies its sta- closely related (see figure 1), but tus as a unique species. not his determination that P. humilis is a subspecies of P. officina- 5) Hong supports his decision lis. to divide section Pæonia into 3 sub-sections with the statement 4) Based on the DNA results, that three sections are “implied” P. mlokosewitschii is a hybrid by the DNA studies of Sang et al. species, which has the same My analysis of Sang’s results in- parentage as five other hybrid dicates that there is very strong species, including P. mascula, justification for dividing section P. clusii, etc. Therefore, these Pæonia into only two sub-sec- six hybrid species form a large tions based on the separate “sister group” based on their evolutionary paths of the two common parentage and must ancient clades within section all be grouped together with the Pæonia. The rationale for this other members of their group. conclusion can best be under- The idea that some in this sis- stood by examining Table 1. ter group have higher or lower species status than others in Considering the complexity of the group seems completely ar- the evolutionary history of sec- tificial. The question of whether tion Pæonia caused by extensive they are species or subspecies natural hybridization, it seems in this group seems like a minor quite remarkable that this com- point. P. banatica, on the other plexity can be captured in the hand, has unique parentage not simple grouping scheme shown shared by any other species and in Table 1, which consists of only thus should clearly be consid- two sub-sections. The philoso- ered to have full species status. phy for constructing this table Its placement has already been was dominated by the single discussed above. P. wittmanni- overriding principal that parents ana is a rare second-generation and their offspring are closely re- hybrid species with four differ- lated, and thus, must be grouped

398-37 Table 1. The proposed grouping of the 19 hybrid species in section Pæonia and the percent contribution which comes from each of their ancient ancestors. together. Applying this philoso- species, all of which are derived phy to section Pæonia, results in from various interclade crosses. only two groups or sub-sections. This group is special because The first sub-section (Group it contains all of the remaining 1) consists of the five ancient species that are carrying the (non-hybrid) species from Clade genes of the two ancient ances- 1 (and includes the extinct spe- tors from Clade 2 that are now cies, ES-C1) and all of the (7) in- extinct, ES1 and ES3 (see Figure traclade hybrid species that are 1). It should also be noted that derived from the various natural all of the members of this group crosses between these (5) an- are of hybrid origin, including cient species. Therefore, Group P. mairei. Thus, this group is a 1 is a pure group with all of its mixed group with all of its mem- members carrying only Clade 1 bers containing genes from both genealogy. The second sub-sec- clades. For the most part, the tion (Group2) is composed of P. genealogy of the hybrid species mairei (the only existing ancient in Group 2 is equally divided be- species from Clade 2) and the tween Clade 1 and Clade 2, with extinct species, ES-C2-2, as well 50% coming from each clade. In as the other remaining hybrid fact, P. wittmanniana is the only

398-38 Figure 3. The Scientist’s Version of the proposed new unified classification for the genus Pæonia based on the recently updated phylogeny from the DNA sequence studies of Sang et al.1,2,3 and the recent taxonomical study of Hong4.

398-39 exception, thus making it a little best characteristic for determin- more difficult to properly classi- ing the taxonomical relation- fy. ships between various species and thus may also explains why In my view, there is no clear the classification of Stern re- need, benefit or justification for mained the standard for such a any further division of this sec- long period of time. As a result, tion. Thus, sub-section Group 1 I have arranged the species and relates only to Clade 1, where- sister groups of species in each as sub-section Group 2 relates sub-section approximately ac- equally to both Clade 1 and cording to their degree of foli- Clade 2). These relationships age dissection, with the species are immediately obvious from exhibiting the highest degree of Table 1. This table also provides foliage dissection at the top and the percent contribution of each those with the least dissection at of the seven ancient species to the bottom. each of the nineteen hybrid species in section Pæonia. It is also The only important classifica- important to note that the spe- tion issues remaining at this cies on both axes of this table are time are the proper placement of intentionally arranged according P. russoi and the five new species to increasing degree of foliage recognized by Hong for which dissection, going from right to there is no DNA sequence data left on the horizontal axis and available. For now, I have placed from bottom to top on the verti- these species according to the cal axis. Thus, the same is also groupings and relationships es- true going along the diagonal tablished by Hong. I have high from the lower right corner to confidence that these place- the upper left corner. As a result, ments will be confirmed once the same approximate foliage the DNA sequence data is avail- dissection variation is also seen able for these species. The same in the classification chart which is also true for the recently dis- results in large part directly from covered and newly recognized this table. This classification yellow peony, P. wendelboi, from chart is shown in Figure 3. the mountains of Iran, which seems to be closely related to P. I believe there is a certain sim- mlokosewitschii. plistic elegance to this grouping system which seems to indicate PART 2 in the Fall Bulletin that foliage type (degree of dissection) is probably the single

398-40 Minnesota Peony Society www.mnpeony.org – Todd O’Halloran - President Good day Fellow APS Members! Here in Minnesota, we have recently seen some sure signs of spring: warm- ing temperatures, melting snow replaced by rains and even a tornado watch here in Southeastern Minnesota (on the same day it was snowing in North- ern Minnesota.) They say if you don't like the weather we're having, just wait a few hours, or apparently just drive a few hours away! We are disappointed that the APS Convention won't be held in-person in Minneapolis, but we are excited for the upcoming plans for online meetings. We also cancelled our usual Winter Educational meeting, and so far are tak- ing a "wait and see" attitude toward the near future, but are hopeful that, as the Governor continues to lift restrictions , we will be able to have an in-person flower show and exhibition this June. Fingers crossed! We will be having a Board of Directors meeting shortly to discuss our plans, tentative as they may be. We wish you all a beautiful, warm and early spring in your gardens and lots of beautiful peony blooms! Looking forward to seeing you online soon!

Duluth Peony Society Elizabeth Donley - President Hello from Duluth: sunny and snow covered, as we got 6 new inches last week. We are looking forward to returning to some of our usual peony cele- brations and neighborhood garden tours (maybe even, weather permitting, an outdoor meeting. Masked still. ) Our Leif Erickson Park garden, overlooking Lake Superior, was spectacular last season. You are all welcome to stop if you are headed up the north shore the end of June.

398-41 Wisconsin Peony Society www.wipeonysociety.org Cory Tischman, President – contact at [email protected] With the early warm-up here in Wisconsin it’s hard not to think about peonies emerging from the ground. I have to stop myself from digging around and checking to see if they made it, as they usually are fi ne and I just end up break- ing off emerging buds. It’s hard to narrow down my favorite peonies as each one brings its own beauty and personality. Some of the cultivars that I am excited to see emerge is the ever handsome foliage and fl ower of 'Nosegay', the stout and punchy red color of 'Burma Ruby', the mesmerizing color combination and fl ower form of 'Perfection', and the sweet lemon fragrance from 'Bartzella'. We recently had our spring educational meeting and biennial elections. We have a great crop of new members to the Board of Directors, and I want to give a special thanks to Jeff Delaney, Helen Finazzo, and Tracey Nelson, for the years and dedication as Treasurer, Newsletter Editor, and Recording Secretary, respectively. For our educational portion, we had an excellent online Q&A panel with notable peony experts: Don Hollingsworth, Harvey Buchite, and Nate Bremer, with the special guest appearance by Martin Page! It was a rare treat Durable, affordable & made in the USA! to have all these people in one place to • Now with rounded corners • share their knowledge and experience with peonies. We are fi nalizing our upcoming events for members and will be announcing the details in our newsletter and website. If you’re interested to volunteer or be in a leadership position in one of our committees, you can contact me using the email address above. Most of the positions can be done remotely. If you INTRODUCTORY OFFER want to fi nd out about what we are do- Sample pack of 8 styles ing here in Wisconsin, check out our plus a marking pencil website or visit us on Facebook: ‘Wis- just $7.50 consin Peony Society Group’. EverlastLabel.com

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398-44 can definitely be an exciting solo endeavor. I am looking forward most to using 'Nosegay' as a pod parent. It offers interesting genetics, being a second-generation offspring from the original cross of two species, P. tenuifolia and P. daurica subsp. mlokosewitschii (or just P. mlokosewitschii to some people). Last year I was able to use some pollen sent by a friend down South from later blooming hybrids, and I had 37 good seeds from 'Nosegay' x 'Pastelegance'. Time will tell if any of them look good! Maybe if you get bored this spring not hanging out with other people, you can make a few crosses. You never know if you might have a winner on your hands.

As always, we are constantly on the lookout for interested individuals who would like to volunteer or be interested in a leadership position on one of our committees. If you are interested, or maybe want to know more, you can contact me using the email address above. Otherwise, if you want to find out about what we are doing here in Wisconsin, check out our website: www.wipeonysociety.org or visit us on Facebook: ‘Wisconsin Peony v Society Group’.

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SEE CONTEST INFORMATION ON THE APS WEBSITE americanpeonysociety.org/news-events/ if-wishes-were-peonies/#virtual-show e wind upon its way whispered the boughs of May, And touched the nodding peony owers to bid them waken. —Siegfried Sassoon