Palmetto Vol. 24(4)

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The Quarterly Journal of the Florida Native Plant Society

Palmetto

Fig.2

Michael Grunwald’s excellent volume is the perfect primer for anyone interested in how the Everglades has been brought to the brink of disaster and for those who hope ardently for its renaissance.

The Swamp lives up to its subtitle – The Everglades, Florida, and the Politics of Paradise. Grunwald begins literally, at the bottom of Florida – exploring the basic geology underlying the Everglades, “the first 300 million years – abridged.” He then examines a key feature of the Everglades environment – water and its majestic slowness as it transits the terrain, quoting an early

The Swamp

The Everglades, Florida, and the Politics of Paradise

explorer – “A raindrop that fell in its headwaters in central Florida could have taken an entire year to dribble down its estuaries at the tip of the peninsula.”

By portraying the shenanigans behind the effort to drain the swamp and make it “useful for society,” Grunwald chronicles the mixture of visionaries and rascals who mingled and contested to finish the swamp once and for all, and the naturalists and unlikely heroes who finally turned the tide toward restoration.

An award winning journalist for the Washington Post, Grunwald began investigating the Everglades story because of his interest in the U.S. Army Corps of Engineers, which he calls “dysfunctional.” His exhaustive collection of sources for each of the 18 chapters are listed in a section of annotated notes at the back of the book, and are evidence of the level of detail he brings to the story.

On the cover – The Pawpaw Quiz

No doubt you’ve seen the colorful and varied flowers arrayed on this month’s PALMETTO cover. Now test your knowledge and see if you can outsmart those tricky and variable pawpaws by guessing their names. Answers are on page 15.

Over one hundred years of mismanagement of the water that ought to flow south to Florida Bay has yielded a seriously crippled Everglades. Its heart is still intact, but for how long? What action is needed to ensure that the Everglades is restored to the best it can be today? That remains to be seen, since politics continues to play an important role in decision making. The story doesn’t end with the close of Michael Grunwald’s book, and visionaries and rascals still struggle over the fate of the Everglades.

Carl Hiassen puts it this way – “The Swamp is the best thing I’ve read about the Everglades. The story of what’s happened to this haunted and magical wilderness has the epic ingredients of a great novel –greed, betrayal, carnage and valor – and Michael Grunwald has beautifully captured it all for history.”

Palmetto is in need of articles on native plant species and related conservation topics, as well as high-quality botanical illustrations and photographs. Contact the editor for guidelines, deadlines and other information at pucpuggy@bellsouth.net, or visit www.fnps.org and follow the links to Publications/Palmetto.

The purpose of the Florida Native Plant Society is to preserve, conserve, and restore the native plants and native plant communities of Florida.

Official definition of native plant: For most purposes, the phrase Florida native plant refers to those species occurring within the state boundaries prior to European contact, according to the best available scientific and historical documentation. More specifically, it includes those species understood as indigenous, occurring in natural associations in habitats that existed prior to significant human impacts and alterations of the landscape.

Organization: Members are organized into regional chapters throughout Florida. Each chapter elects a Chapter Representative who serves as a voting member of the Board of Directors and is responsible for advocating the chapter’s needs and objectives. See www.fnps.org

Board of Directors

Executive Officers

President – Shirley Denton

Past President – Bob Egolf

Vice President,Admin. – Ann Redmond

Vice President,Finance – Travis MacClendon

Treasurer – Susan Thompson

Secretary – Ameé Bailey Speck

Committee Chairs

Communications – Cindy Liberton

Conference – Charlie Pedersen

Conservation – Suzanne Kennedy, Eugene Kelly

Development – JoAnne Trebatoski

Education – Judy Gregoire

Finance – Travis MacClendon

Governmental Policy – Kim Zarillo, Annie Schmidt

Landscape Awards – Mike Kenton

Membership – Mary Ann Bolla, Lynka Woodbury

Public Lands – Charlie Pedersen

Publications – Eileen Szuchy

Science – Paul Schmalzer

Directors-at-large

Lauren Day (2006-2008)

George Kish (2006-2008)

Laurel Schiller (2006-2008)

Sue Dingwell (2007-2009)

Ray Jarrett (2007-2009)

Anne Birch (2007-2009)

To contact board members: FNPS Administrative Services

Phone: (321) 271-6702 info@fnps.org • www.fnps.org

To join or for inquiries: Contact your local Chapter Representative, or call, write, or email FNPS, or visit our website.

Florida Native Plant Society PO Box 278

Melbourne FL 32902-0278

Phone: (321) 271-6702

info@fnps.org • www.fnps.org

Webmaster – Paul Rebmann

Contract Services

Executive Director – Karina Veaudry

Accounting Services – Joslin & Hershkowitz

Administrative Services – Cammie Donaldson

Editor, Palmetto – Marjorie Shropshire

Editor, Sabal Minor – Rosalind Rowe

Palmetto

Features

4The “False Pawpaws” – Deeringothamnus

The question remains – is Deeringothamnus different enough from Asimina to be recognized as a distinct genus? The discovery of a probable hybrid between the two makes the topic even more interesting. Dr. Eliane Norman explores the differences and similarities between Florida’s pawpaws and “false pawpaws.”

8Native Plants are the Stars of the Show

At the annual FNPS Conference, the Landscape Awards Committee honors deserving homeowners, restorationists and professionals for their beautiful and innovative use of native plants in the landscape. See the stars of the 2007 Landscape Awards show and be inspired to show off your own landscape in 2008.

12Does your Pawpaw Smell Flowery?

Scientists are increasingly interested in the role that plant scent compounds play in plant-animal interactions. But floral scents are also becoming a popular topic for gardeners, horticulturists and florists. Katherine Goodrich looks at floral scent in Florida’s pawpaw species.

Departments

2 Book Review

16 FNPS Chapters

ON THE COVER: Flowers of Asimina and Deeringothamnus species. Photos 2, 3, 4, 6, 7, 8 and 9 by Paul Rebmann, www.wildflphoto.com. Photos 1 and 5 by Marjorie Shropshire.

The Palmetto (ISSN 0276-4164) Copyright 2007, Florida Native Plant Society, all rights reserved. No part of the contents of this magazine may be reproduced by any means without written consent of the editor. The Palmetto is published four times a year by the Florida Native Plant Society (FNPS) as a benefit to members. The observations and opinions expressed in attributed columns and articles are those of the respective authors and should not be interpreted as representing the official views of the Florida Native Plant Society or the editor, except where otherwise stated.

Editorial Content: We have a continuing interest in articles on specific native plant species and related conservation topics, as well as high-quality botanical illustrations and photographs. Contact the editor for submittal guidelines, deadlines and other information.

Editor: Marjorie Shropshire, Visual Key Creative, Inc. ● pucpuggy@bellsouth.net ● Telephone (772) 232-1384 ● 855 NE Stokes Terrace, Jensen Beach FL 34957

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Images, top to bottom: Deeringothamnus rugelii – photo by Dr. Eliane Norman Award-winning native plant landscape design – photo by Eugene Stoccardo. Asimina incana – photo by Katherine Goodrich. Top right: Asimina obovata –drawing by Marjorie Shropshire.

The “False Pawpaws”

History, Biology and Conservation of Deeringothamnus

John Kunkel Small, a pioneer explorer of the southeast, especially Florida, started his botanical explorations to the southern states as early as 1891, before his appointment as curator at the New York Botanical Garden. For the next four decades, he spent some weeks or sometimes months of most years in the south. Fortunately for him (and for us) he was able to get sponsors for this travel, as he never learned to drive. Thus on April 28, 1923, while he was botanizing with his friends, C. A. Mosier and J. DeWinkler in the ”uninhabited pineland wilderness between Punta Gorda and Fort Myers,” Charlotte County – an odor that wafted in the air alerted him to the fact that he was about to encounter a new plant, even before he had an opportunity to examine it. He went on to name (1924) this plant Deeringothamnus pulchellus, meaning “beautiful little Deering’s shrub,” quite a mouthful. Charles Deering was Small’s friend and patron, son of the founder of the Deering Harvesting Machine Co. and International Harvester, who owned a large estate south of Miami. Small mentioned that Deeringothamnus resembled Asimina, but differed from it in having narrow petals, all the same size, with no floral chamber around the sex organs and with a flat receptacle.

Deeringothamnus pulchellus – photo by Kate Goodrich

A few years later, when he was exploring the east coast of Florida, Small came upon another species of this genus. This time the flowers were smaller, yellow and lacked the scent that had lead to the discovery of its congener. However this species had already been discovered in May 1848 by a German pharmacist-botanist, Ferdinand Rugel. This collection was sent to his English sponsor, J. B. Shuttleworth and laid there until the American botanist, B. L. Robinson studied and named it Asimina rugelii. Small (1930) made the new combination, D. rugelii (B. L. Rob.) Small.

The question remains, is Deeringothamnus different enough from Asimina to be recognized as a distinct genus? In the past some have said yes, such as Kral (1960), who wrote his thesis on the two genera and Fries (1939, 1959), a world-wide authority on the family Annonaceae. Others were in favor of lumping them together (Wood 1958, Wilbur 1970). It should be pointed out that none of these authors were aware of the existence of hybrids between the two genera. We have seen three plants that most likely reflect this mixed parentage. One was a potential hybrid of D. rugelii and A. pygmea and the other two of D. pulchellus and A. reticulata. [Fig. 1] Details are given in Norman, 2003. The main morphological differences between the two genera are listed in the Table and will be commented on

briefly. Certainly at the morphological level, the two are quite distinct but it may be that molecular studies will not support these distinctions (Abbot and Neubig, per. com.).

The presence of stomates on both leaf surfaces is an unusual situation found in almost no other member of this family. In other groups this feature has been associated with xeric conditions which may have prevailed when Deeringothamnus evolved. The diminutive stature may have been associated with the fire regime present during its development. The small flowers with less specialized petals accompanied by fewer sex organs, especially stamens, are traits difficult to explain. The much lower pollen to ovule ratio indicates a trend to more self-pollination. This characteristic is accompanied by a very low level of pollinators.

TABLE AsiminaDeeringothamnus

Differences Between Asimina and Deeringothamnus

HABIT

Shrubs-treesDwarf shrubs

LEAF EPIDERMISLower surface onlyBoth surfaces Stomates

BRACTSTwo-oneNone

FLOWER SIZEMedium-largeSmall

PETALS

ShapeTwo distinct seriesAll the same SculpturingOn inner petalsNone

ChamberYesNo

RECEPTACLEHemisphericFlat

STAMENS

Number 70-20016-34

Connective (tip)Knob-likeTongue-like

POLLEN

Perforations4-5 µmUp to 9-12 µm

To ovule ratio1240 - 3200 : 1325 - 413 : 1

POLLINATORSPrimarily beetlesVery infrequent (flies, thrips, beetles)

FIG. 1.
A. Asimina reticulata. B. putative hybrid. C. Deeringothamnus pulchellus. Photo by Ed Norman enhanced by John Weishampel. (from Taxon 2003)

Both species of Deeringothamnus are rare, and are listed as federally endangered (USFWS 1886, Coile 2000). Ward (2001), without critical study, concluded that there are intermediates between the two species of Deeringothamnus and based on this erroneous supposition, combined them into one species with two varieties. This is doubly unfortunate, since these new combinations were adopted by Wunderlin and Hansen (2003).

Deeringothamnus rugelii, known commonly as Rugel’s pawpaw or yellow squirrel banana, has never been found outside a narrow corridor in eastern Volusia County from the Lake Ashby area to Tiger Bay State Forest. Approximately 15 populations are known and a fourth of these are on public lands. Most populations occur on Immokalee fine sands which are poorly drained soils with some organic matter in the upper 5-8 inches. They grow in flatwoods with an open canopy of longleaf or slash pine and an understory of wiregrass and shiny blueberry, dwarf huckleberry, common pawpaw and saw palmetto.

Typically the plants die back each winter. In some cases plants may overwinter but then their woody branches tend to produce only leaves the following season. Flowers are produced chiefly on new shoots which develop from the upper portion of thick tap roots. Rugel’s pawpaw flowers from midApril to early June. In a population which has not been recently disturbed by fire or mowing, only 3-15% of plants can be expected to flower in a particular year. If the population is disturbed, many of the plants can be expected to bloom within six weeks of the disturbance (Helkowski & Norman 1997). The nodding flowers mature from the base upward, and have six or more linear subequal fleshy yellow petals in two whorls. [Fig. 2A] Rarely, plants with purple-red flowers are found. [Fig. 2B] As in other members of the custard apple family, the female organs mature first. This can be observed when the one to four stigmas glisten with a sticky exudate. Then the 15-25 short-stalked stamens turn whitish-grey and dehisce revealing (with a hand lens) small worm-like columns of pollen tetrads. There is often an overlap of at least one day between the two sexual phases (Norman 2003). Fruiting is rare; in several hundred flowers studied in two separate years, less than 2% set fruits (Norman 2003). The fruit is made up of one or

FIG. 2.
A. Deeringothamnus rugelii. B. red form of D. rugelii. C. developing fruits of D. rugelii. D. D. pulchellus. Photos A and D by Steve Shirah, B and C by Eliane Norman. (from Taxon 2003)

more yellowish-green, smooth-skinned, peanut-like structures, each with one to several large brown seeds embedded in a custard-like tissue. [Fig. 2C)]

Deeringothamnus pulchellus, beautiful pawpaw has a wider distribution than its congener. It is known from Charlotte and Lee Counties in southwest Florida as well as from a few populations in eastern Orange County, a hundred and fifty miles away. A third of the 27 populations known from the west coast are on public land (Johnson 1999), none from the east coast is protected. The beautiful pawpaw occurs in poorly drained sandy soils, in flatwoods. In Orange County it is restricted to Pomello fine sand, while in Charlotte and Lee Counties, the largest number of populations grow in Wabasso soils. This soil has a clayey loam texture at a depth of over two feet (Johnson 1999). The associates of this species are similar to those of D. rugelii, namely slash or long leaf pine, wiregrass, saw palmetto, dwarf live oak, wax myrtle, common pawpaw and hatpins.

The beautiful pawpaw is a more highly branched shrub than Rugel’s pawpaw. It blooms from mid-March to early May, and also reacts to fire by flowering. It differs from Rugel’s pawpaw by having longer pedicels, longer, thinner sepals, and thinner and longer reflexed petals – often more than six. [Fig. 2D] Its very sweet scent is somewhat like confederate jasmine, an odor not found in other pawpaws, even ones with white flowers. The sex organs are similar, and there may be an overlap in the maturation of the reproductive organs. In a study of almost 500 flowers in a particular year, only 1.2% set mature fruits (Norman 2003). The fruits and seeds are similar to those of D. rugelii.

The fruits of Deeringothamnus take three to four months to mature. Over the years, we have seen a considerable number of dried seeds or fruits scattered around the base of plants of both species. When the fruit is mature, the seeds have a minute immature embryo and a thick seed coat. Even when planted under “good conditions,” (in an area where Deeringothamnus are already growing), it will take between six months to a year before they show any growth above ground (Norman 2003).

At least some of the reasons for the rarity of both species of Deeringothamnus are apparent: plants do not flower readily unless disturbed, the flowers are small and produce small amounts of pollen, they attract few pollinators and they have low fruit and seed production. The seeds need to be buried and kept reasonably moist for the embryos to grow and germination to occur. However once established the plants probably survive for several decades, if given enough living space. What can be done to keep these species from becoming extinct?

Ultimately, both species of Deeringothamnus can only be preserved through acquisition and proper management of their habitats. We need to learn more about maintaining and enhancing the populations. If fire is not feasible, roller chopping or mowing may be beneficial in suppressing competition and enhancing flower production (Helkowski and Norman 1997). Our experiments indicate that hand cross pollination enhances fruit and seed production (Norman 2003), but we also found that such seeds germinated only half as often as seeds from open-pollinated flowers (Norman 3003). Transplanting is a difficult process as Deeringothamnus have a large tap root. [Fig. 3] Plants must be removed quickly and not allowed to dry out before replanting. Success requires enough rain (but not too much), so plants can become established. Most successes in transplanting and seed germination have been obtained when Deeringothamnus are planted in a field

3.

Digging up Deeringothamnus pulchellus from a threatened area. Photo by Sandy Carnival.

FIG.

FNPS 2007 Landscape

Each year at the FNPS Annual Conference, the Landscape Awards Committee presents “Design with Natives,”a program that salutes the use of native plants in both restorative and traditional landscapes. Projects from around the state are submitted in a variety of categories, and the best are chosen to receive an award. Both professionals and non-professionals are invited to enter their work for consideration. From complete site restorations to tiny butterfly gardens in a local schoolyard, “Design with Natives” provides everyone an opportunity to be honored for their native planting efforts. Additional photos of this year’s winners and an application for the 2008 awards may be found on www.fnps.org. Go to “Programs” and follow the link to “Awards and Grants.”

Ecosystem Restoration Landscape – First Place

Coe Visitor Center, Everglades National Park West of Homestead and Florida City.

Dade Chapter – Florida Native Plant Society, Designer, Contractor, Maintenance. Everglades National Park, Owner.

Plantings were selected to mimic the nearby Everglades hammock ecosystem of pine rocklands and wetland habitats, and to enhance the views from the Coe Visitor Center.

Awards

Many challenges and site constraints were overcome to create the wildflower and butterfly garden at the Enchanted Forest Sanctuary environmental center. The primary goal of the design was to provide the maximum number and variety of pollinator plants (including both host and nectar plants) while demonstrating a holistic approach to gardening.

A diverse range of habitat was created, including space for wetland species such as the beautiful purple pickerelweed (Pontederia cordata).

The walking path narrows periodically to let visitors experience plants such as the coneflower (Echinacea purpurea) up close. Plants immediately adjacent to the pathways were specifically chosen for one or more of the following qualities: height, scent, color, texture, and durability.

The Enchanted Forest Sanctuary Butterfly/Pollinator Garden Titusville, Florida. The Friends of The Enchanted Forest, Designer. Keep Brevard Beautiful and The Friends of The Enchanted Forest, Contractor. The Enchanted Forest Sanctuary, Owner. “Deadheaders” Garden Volunteers, Maintenance.

Wildflower & Butterfly Garden Landscape – First Place

FNPS 2007 Landscape Awards

Homeowner Landscape – First Place

Before designing their landscape, Eugene and Cathy Stoccardo studied historical records to find the soil type and habitat that originally occurred on their home site.

Two years after installation, birds, butterflies, and insects visit this well-planned landscape, showing that even an urban, residential yard can be transformed into a native plant oasis.

Eugene & Catherine Stoccardo Residence, Orlando, Florida. Eugene Stoccardo, Catherine Stoccardo, Designer, Owner & Contractor.

Ticknor Residence

Santa Rosa Beach, Florida.

David A. Ilfrey, Jr. NativeDave.com, Designer. Angie & Steve Ticknor, Owner.

An example was set for responsible land stewardship while appealing to traditional community standards. This sustainable landscape design conserves water, energy, time and money and restores eco-regional identity.

Professional Residential Landscape – First Place

Ugarte & Associates, Inc., Palmetto, Florida

Allen Garner & Associates, Designer. Charlie & Patsy Ugarte, Owner. Tom Heitzman, Sweetbay Nursery, Contractor.

This beautiful use of native plant color and texture has set a precedent in the area and others have inquired about converting their traditional lawn oriented landscapes to natives.

Additional FNPS 2007 Landscape Awards

Ecosystem Restoration Landscape – Second Place

Felts Audubon Preserve Restoration Palmetto, Florida.

Manatee County Audubon Society and Florida Department of Environmental Protection, Designer, Contractor & Maintenance. Manatee County Audubon Society, Owner.

Homeowner Landscape – Second Place

Marilyn Smullen Residence New Port Richey, Florida. Marilyn Smullen, Designer, Owner.

Ecosystem Restoration Landscape – Third Place

Circle B-Bar Reserve

Polk County Environmental Lands Program, Polk County. Laura Namm NeSmith, Water & Air Research, Inc., Designer. Water & Air Research, Contractor.

Polk County Board of County Commissioners, Owner. Polk County Environmental Lands Program, Maintenance.

Homeowner Landscape – Third Place

Sharon Holding Residence

New Port Richey, Florida.

Sharon Holding, Designer, Owner

Commercial Landscape – First Place

Does your pawpaw smell flowery or fermented?

Communicating attributes of floral scent can be difficult. Most people are familiar with the pleasing floral scent of a rose, a gardenia or a magnolia flower. However, imagine trying to communicate the difference between these floral scents. Words such as “sweet” or “pleasant” describe them all, but it is more difficult to find words which describe the unique qualities of each one. The perfume and wine industries employ researchers and technicians who have fine-tuned their scent perception and developed a detailed vocabulary for scent. For those of us outside of these industries, our scent perception and vocabulary is markedly less refined.

Until recently, the scientific study of scent was also relatively confined to the perfume or wine industries. Advances on scent sampling and analysis techniques have broadened the scope of scent study, and plant biologists, insect physiologists and ecologists are increasingly interested in the role that plant scent compounds –especially floral scent compounds – play in plant-animal interactions. Floral scents are also increasingly popular among gardeners, horticulturalists, and florists (Imato, 2005).

About the Author

Katherine Goodrich’s scent study of Asimina triloba expanded to cover the floral display (including floral scent) of all species of Asimina and Deeringothamnus, as well as their close tropical relative, Annona glabra. Kate says “The floral display of these plants are of interest to me because they represent extra-tropical genera from a predominantly tropical family (Annonaceae). I am interested in their evolutionary history as they diverged from their closest tropical relatives.” Kate is entering her last year of the Ph.D. program at the University of South Carolina, and will graduate in December, 2007.

My own scent research focuses on the genera Asimina and Deeringothamnus (Annonaceae), commonly know as pawpaws. Pawpaws present an interesting case study in floral scent. Half the species of Asimina have flowers which emit “flowery” scents used frequently in the perfume industry. The other half of Asimina species have flowers which emit “fermented” odors – scents produced by microbes during the fermentation process – which have been extensively studied by the wine industry. Many of the scent compounds emitted by pawpaw flowers are well documented and easily identifiable. The two species of Deeringothamnus have scents which differ from Asimina species and from each other. D. pulchellus has a distinctively strong, pleasant scent, while D. rugelii has a faint, unpleasant aroma.

Four of the eight Asimina species have white flowers and are limited to Florida and southern Georgia and Alabama

FIGURE 1
A. Asimina reticulata. B. Asimina incana. C. Asimina angustifolia. D. Asimina obovata
Photos by Katherine Goodrich.

(A. reticulata, A. angustifolia, A. incana and A. obovata). [Fig. 1] The other four species of Asimina have small maroon or maroon and white flowers; two maroon-flowered species occur in Florida (A. tetramera and A. pygmea), while the other two have larger ranges extending farther north across the eastern U.S. (A. triloba and A. parviflora) (Kral, 1960). [Fig. 2]

The floral scent of Asimina species has an interesting correlation to floral pigmentation. Species with maroon floral pigmentation all share yeasty “fermentation” floral odors, while species with white or cream-colored petals have more varied “sweet”, “soapy/waxy” or “perfumy” floral scents. The reason for this difference is not clearly understood, but it seems that the floral scent of Asimina species is integrally tied to the overall floral phenotype of each species.

Chemical analyses show that many of the scent compounds produced by maroon-flowered species of Asimina are the same as those produced by active Baker’s yeast (Goodrich et al., 2006; Goodrich & Raguso unpub. data). These fermentation compounds give the flowers a scent similar to red wine or rising bread dough. Although these flowers and Baker’s yeast both produce ethanol, ethyl acetate, methyl butanol and hydroxy-butanone, these compounds are produced in different ratios by the flowers than they are by yeast. However, to human perception the scents are very similar.

Most Floridian species of Asimina have large white, droopy flowers and are a common sight in pastures and along frequently-mowed roadsides. The scent of these white-flowered species is typically described as “pleasant,” although it varies more between species than the yeasty scent of maroon-flowered Asimina A. reticulata has, in my opinion, the most pleasant floral scent of the genus. It has a sweet, almost fruity odor. Asimina incana, on the other

hand – has a heavy sweet odor with an acrid, soapy note which at high concentrations is almost unpleasant. Chemically, all of the white-flowered species share hydrocarbons and several terpenoid compounds in their floral scent (Goodrich & Raguso, unpub. data).

Three of the four white-flowered species share several aldehydes and alcohols typically found in the scent of lilacs. Other pleasant-smelling compounds found in some whiteflowered species include benzaldehyde (pleasant, almond-like odor) and phenyl ethyl alcohol.

The floral phenotypes of Deeringothamnus species differ from one another and from species of Asimina (Norman, 2003). Both species of Deeringothamnus have floral

FIGURE 2
A. Asimina tetramera. B. Asimina parviflora. C. Asimina pygmea. D. Asimina triloba
Photos A, B, C by Katherine Goodrich. Photo D by Paul Rebmann, www.wildflphoto.com
FIGURE 3
Deeringothamnus pulchellus. Photo by Paul Rebmann, www.wildflphoto.com

Does your pawpaw smell flowery

or fermented?

Deeringothamnus pulchellus, the beautiful pawpaw, has flowers which are small, white and have a strong, pleasant scent. The scent is dominated by compounds such as benzaldehyde, linalool, methyl benzoate and benzenacetaldehyde which are also found in some whiteflowered Asimina species. The floral scent of D. pulchellus also contains ethyl benzoate and veratrol not found in any Asimina species. [Table 1]

innate response from a naive insect. Of course, the floral scents of these genera are also likely influenced by factors outside of pollinator selection pressures. Evolutionary history, phylogenetic constraints, neutral genetic variation, and plant nutrition may all play a role in the floral scent produced by each flower.

A great deal of research remains to be done on the floral displays of Asimina and Deeringothamnus, but floral scent is clearly an important part of this display. A study of floral scent within these genera provides an interesting context in which to ask questions of pollinator attraction, floral evolution and plant biochemistry. As studies of floral scent become more widely published and discussed, our

Table 1: Distinctive scent compounds found in Asimina and Deeringothamnus Compounds listed represent major and/or distinguishing components of the floral scent for each species of Asimina and Deeringothamnus. Descriptions of scent perception are from personal experience of the author or taken from descriptions provided on flavornet.org.

Asimina species

Deeringothamnus species

DISTINCTIVE CHEMICAL maroon or maroon/white flowerswhite flowers COMPOUNDHUMAN PERCEPTIONTRIPARPYGTETRETINCANGOBOPULRUG FERMENTATION PRODUCTS

ethanolsweet, pleasant, solvent*

methyl butanolwhiskey, malt, burnt*

hydroxybutanonebutter, cream*

butanediolonion*

ethyl acetatepineapple*

ALIPHATIC COMPOUNDS

n-tridecanewaxy

n-pentadecanewaxy

heptadecadienewaxy

AROMATIC COMPOUNDS

Deeringothamnus rugelii, Rugel’s pawpaw, has small, yellow flowers with a very slight, unpleasant odor. Although to my perception D. rugelii smells slightly similar to the maroon-flowered species of Asimina, it contains ethyl acetate and no other fermentation compounds identifiedin Asimina species. It also contains a number of “unknown” compounds that I am currently working to identify, not found in other species of Asimina or Deeringothamnus. Within Asimina, the small maroon flowers with yeasty scents may represent mimics of food substrates or brood sites for pollinating beetles and/or flies.The alcohols, ketones and aldehydes found in D. pulchellus and many whiteflowered Asimina species may mimic the scents of ripe fruit (prior to more unpleasant fermentation and decay odors). These scents may also mimic food substrates of beetles. It is also possible in each of these cases that the odors produced generate a learned response from pollinating insects. Certain insects may learn to identify particular odor compounds with nutritious floral tissues although the odors themselves may notgenerate an

benzaldehydepleasant, almond-like

phenyl ethyl alcoholsimilar to hyacinth or rose

methyl benzoate prune, lettuce, herb, sweet*

ethyl benzoateflowery, fruity, celery-like*

benzenacetaldehydesweet, perfumy odor

veratrolsweet, floral odor

TERPENOID COMPOUNDS

lilac alcohols and aldehydes flowery, sweet, herbaceous

linaloolspicy floral scent, lavender*

ocimeneherb, citrus, flower*

limonenelemon, orange

caryophyllenesharp, herb

1,8-cineolmint, sweet*

* Source: Flavornet by Terry Acree & Heinrich Arn, http://www.flavornet.org, © Datu Inc., 2004

vocabulary for scent characterization grows, and readers will become more familiar with the language of scent. I hope to help promote this increased awareness of floral scent since it is truly an underappreciated – and integrally important aspect of floral display.

REFERENCES

Acree, T. & H. Arn. 2004. Flavornet. http://www.flavornet.org, © Datu Inc.

Amato I. 2005. Save the flowers: Would-be scent engineers aim to resurrect lost floral fragrance. Science News. 168 (13): 202. http://www.sciencenews.org/articles/20050924/bob10.asp

Goodrich, K. R., Zjhra, M. L., Ley, C. A. & Raguso, R. A. 2006. When flowers smell fermented: the chemistry and ontogeny of yeasty floral scent in pawpaw (Asimina triloba: Annonaceae). International Journal of Plant Sciences. 167(1): 33-46. Kral, R. 1960. A revision of Asimina and Deeringothamnus. Brittonia. 12(4): 233-278. Norman, E. M. 2003. Reproductive biology of Deeringothamnus rugelii and D. pulchellus (Annonaceae). Taxon. 52: 547-555.

Answers: The Pawpaw Quiz

1. Asimina tetramera

2. Asimina parviflora

3. Deeringothamnus rugelii

4. Asimina reticulata

5.Probable hybrid between

A. tetramera and A.reticulata

6. Deeringothamnuspulchellus

7. Asiminaincana

8. Asiminatriloba

9. Asimina obovata

The “False Pawpaws”– History, Biology and Conservation of Deeringothamnus

which already has the particular species growing in it (Workman, per. com., Norman per. obs.). Whether this is due to a mycorrhizal association is not known, nor is anything known about the genetic variability in the two species. Clearly, much remains to be learned about these fascinating plants.

REFERENCES

Coile, N. C. 2000. Notes on Florida’s Endangered and Threatened Plants. Botany Contribution no.38, ed. 3. Dept. Agr. and Cons. Serv. Div. Plant Industry, Gainesville, FL. Fries, R. E. 1939. Revision der Arten einiger Annonaceen-Gattung. V. Acta Horti Berg. 12: 546-554.

Fries, R. E. 1959. Annonaceae. Pp.1-171 in: Engler, A. and K. Prantl (eds) Die naturlichen Pflanzenfamilien, ed. 2, vol. 17aII. Duncker and Humblot, Berlin. Helkowski, J. H. and E. M. Norman. 1997. Effects of fire on an endangered Florida plant, Deeringothamnus rugelii. Florida Sci. 60: 118-123. Johnson, A.F. 1999. a survey for beautiful pawpaw (Deeringothamnus pulchellus) on Babcock-Webb WMA and Charlotte Harbor State Buffer Preserve, Charlotte and Lee counties, Florida. An FNAI Inventory. Florida Natural Areas Inventory. Tallahassee, FL.

Kral, R. 1960. A revision of Asimina and Deeringothamnus (Annonaceae). Brittonia 12: 233-278.

Norman, E. M. 2003. Reproductive biology of Deeringothamnus rugelii and D.pulchellus. Taxon 52: 547-555.

Small, J. K. 1924. Plant novelties from Florida. Bull. Torrey Bot. Club 51: 390.

Small, J.K. 1930. Deeringothamnus rugelii. Addisonia 15:17-18.

U.S. Fish and Wildlife Service (USFWS). 1986. Endangered and threatened wildlife and plants; endangered or threatened status for three Florida plants. Federal register 51: 34415-34420.

Ward., D. B. 2001. New combinations in the Florida flora. Novon 11: 360-365.

Wilbur, R. L. 1970. Taxonomic and nomenclatural observations on the eastern North American genus Asimina (Annonaceae). J. Elisha Mitchell Sci. Soc. 86: 88-96.

Wood, C. E. Jr. 1958. The genera of the woody Ranales in the southeastern United States. J. Arnold. Arbor. 39: 296-346.

Wunderlin, R. P. and B. Hansen. 2003. Vascular plants of Florida. 2 ed. U. Press of Florida, Gainesville.

About the Author

Eliane M. Norman is professor emerita of biology at Stetson University, Deland, Florida, and is the co-author of Andre´ Michaux in Florida

FNPS Chapters & Representatives

For chapter contact information, please visit http://www.fnps.org/pages/chapters/chaptermap.php

1.Broward .........................Molly Taylor ....................................................mollytaylor_piesplants@yahoo.com

2.Citrus ..............................Teddi Bierly......................................................................jbierly@tampabay.rr.com

3.Coccoloba ......................Dick Workman................................................................wworkmandick@aol.com

4.Cocoplum .......................Joan Bausch.....................................................................................jclb@gate.net

5.Conradina .......................Renee Shaffer.................................................................shaffer_m@bellsouth.net

6.Dade ...............................Lynka Woodbury....................................................lwoodbury@fairchildgarden.org

7.Eugenia ..........................Sue Thompson....................................................................sue@pa-services.com

8.Heartland .......................Amee Bailey ..............................................................ameebailey@polk-county.net

9.Hernando ........................Jim Clayton......................................................................geoska10@hotmail.com

10.Ixia .................................Jake Ingram................................................................jakeingramla@comcast.net

11.Keys ................................Lynka Woodbury....................................................lwoodbury@fairchildgarden.org

12.Lake Beautyberry ..........Ronald Plakke............................................................................flplakke@aol.com

13.Lakelas Mint ..................Royce Sisson.................................................................roycesisson@hotmail.com

14.Live Oak .........................Hal Rhodes................................................................rhodeslesstrav2@yahoo.com

15.Longleaf Pine .................Amy Hines.................................................................amy@sidestreamsports.com

16.Lyonia ..............................Ray Jarrett..............................................................................sjarrett1@cfl.rr.com

17.Magnolia ........................Nona Elder.......................................................................nonaelder@pipeline.com

18.Mangrove .......................John & Christine Holyland .....................................christineholyland@comcast.net

19.Naples ............................Ron Echols...................................................................preservecaptains@aol.com

20.Nature Coast ..................Susan Vaughn..........................................................................vaughn_s@firn.edu

21.Palm Beach ....................Ray Miller................................................................................rmillerj@sfwmd.gov

22.Pawpaw .........................Paul Rebmann..................................................................wildflpaul@earthlink.net

23.Paynes Prairie ................Erick Smith..................................................................................edsmith@ufl.edu

24.Pinellas ..........................Debbie Chayet & Ray Wunderlich III.....dchayet@verizon.net; rayrunner@yahoo.com

25.Sarracenia ......................Lee Norris .................................................................................lnorris@ectinc.com

26.Sea Rocket ....................Paula Berntson.......................................................................jepasm.3@juno.com

27.Serenoa ..........................Nancy Armstrong..............................................................pinelilywild@yahoo.com

28.South Ridge ...................Stacy Smith...............................................................ssmith@archbold-station.org

29.Sumter ............................H. Tuxbury..............................................................................HTuxbury@cfl.rr.com

30.Suncoast .......................James Wheeler............................................................jamesh-wheeler@msn.com

31.Tarflower ........................Rick Ehle........................................................................................(407)349-0502

The Florida Native Plant Society PO Box 278 Melbourne FL 32902-0278

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Please contact: Travis MacClendon, Vice President of Finance, FNPS Blountstown, Fl 32424 850.674.4408 travismac@wfeca.net

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Palmetto Vol. 24(4) by Florida Native Plant Society - Issuu