JARS v47n3 - Exploring the Deciduous Azaleas and Elepidote Rhododendrons of the Midwestern United States

Exploring the Deciduous Azaleas and Elepidote Rhododendrons of the Midwestern United States
Mark P. Widrlechner 1 , Richard A. Larson 2 , Sharon K. Dragula 1

Introduction
A great diversity of deciduous azalea cultivars and many elepidote rhododendron cultivars include native American species in their parentage. Many collections from wild populations of native species have been made for years from states along our nation's eastern seaboard. More recently, McCullough (1985) and Boge (1986, 1987, 1988) have reported on extensive collections made along the West Coast.
But wild azalea and rhododendron populations in the Midwest have received little notice. Azaleas and rhododendrons are generally not common in the region and can easily be overlooked. One of the most ambitious collectors of native azaleas and rhododendrons, Henry Skinner (1955), skipped over midwestern sites during his travels to areas in which these plants are more abundant.
Three azalea species, the roseshell azalea ( Rhododendron prinophyllum [Small] Millais), the pinxterbloom azalea ( Rhododendron periclymenoides [Michx.] Shinners), and the flame azalea ( Rhododendron calendulaceum [Michx.] Torrey), and the rosebay rhododendron ( Rhododendron maximum L.) reach the northwest edge of their native range in isolated populations stretching from the Ohio River near Wheeling, West Virginia to south-central Missouri. At such sites, January mean temperatures range from 30 to 34°F, about the same as those found from New York City south to Wilmington, Delaware; but in July, mean temperatures are similar to those farther south, such as Washington, DC and Winston-Salem, North Carolina (75 to 78°F). Although azaleas and rhododendrons from midwestern states should be better adapted to climatic extremes than those populations found farther east, they are not normally available to hybridizers or nurseries.
In 1989, we received a grant from the Research Foundation of the American Rhododendron Society to investigate native midwestern species, with the goal of collecting pollen and seed samples for the ARS Pollen Bank and Seed Exchange, and also to analyze soils to see if we could locate plants thriving on sites with soils less acidic than typical. During 1990 and 1991, with the help and advice of many foresters and botanists, we were able to visit many sites in Ohio, Missouri, and Illinois. These trips led to the collection of pollen from 16 populations in Ohio, 11 in Missouri, and 3 in Illinois. A similar number of seed collections was made, including all four species native to the region. In addition, 33 soil samples were taken for analysis of pH, phosphorous, potassium, and organic matter. Collection sites are mapped on Figure 1.

Map of Rhododendron collection sites.
Figure 1. Map of Rhododendron collection sites. Legend: C = Rhododendron calendulaceum ;
M = Rhododendron maximum ; P = Rhododendron prinophyllum ; X = Rhododendron periclymenoides .
Note: Nearby sites were combined to eliminate overlapping symbols.

We will summarize our experiences in three sections: Rhododendron prinophyllum in Missouri and Illinois; deciduous azaleas in Ohio; and Rhododendron maximum in Ohio.

Rhododendron prinophyllum in Missouri and Illinois
Two of us (Widrlechner and Dragula) travelled to the southernmost part of Illinois during the second week of May in 1990 and then proceeded west as far as Douglas County, Missouri during the next four days, visiting azalea populations we identified from herbarium records and interviews with foresters and naturalists. We collected pollen, soil samples, and field notes from 14 different populations of the roseshell azalea, R. prinophyllum .
In Missouri and Illinois, R. prinophyllum is usually found on acidic, rocky soils in the understory of oak and oak-shortleaf pine forests. Common companion plants include small red maples ( Acer rubrum L.), sour gum ( Nyssa sylvatica Marsh.), flowering dogwood ( Cornus florida L.), farkleberry ( Vaccinium arboreum Marsh.), and the low-bush blueberry, Vaccinium vacillans Torrey. The azaleas were often growing on north or west-facing slopes and generally did not flower well except along the edges of trails or where the forest was thin.
Flowering characteristics (time of bloom, flower color, and flower size) were rather uniform among the 14 populations we visited in the spring. Throughout this area, the peak flowering occurred between the 10th and the 17th of May. Petal colors only included shades of pink (Royal Horticultural Society [RHS] Color Classes 55, 56, 62, and 65). Tube colors were even less variable (RHS Color Classes 55 and 62, with most 55A). We turned up no true whites, although at least one was described from Sainte Genevieve County, Missouri (Steyermark, 1963) and we heard rumors of others. The palest flowers were found in Ripley County, Missouri (RHS 56C) and the most intensely colored flowers came from a variable population at the Spiva Azalea Park (see photo) east of Fredericktown in Madison County, Missouri (RHS 65C to 65A). Nearly all the flowers lacked petal blotches. Such blotches are not typical of R. prinophyllum , but we did find some plants with faint orange petal blotches at the Spiva Azalea Park. Among all populations, flower diameter, measured with petals spread, varied from 4.75 to 6.0 cm with most between 5.0 and 5.5 cm.

Rhododendron prinophyllum at the Spiva 
Azalea Park
Rhododendron prinophyllum at the Spiva
Azalea Park, Madison County, Missouri.
Photo by Sharon K. Dragula

We found one of the most attractive azaleas of the trip in Oregon County, Missouri in a unit of the Mark Twain National Forest. The plant was very floriferous with clear pink flowers. It stood about a meter tall with a spreading form. A single-plant collection was made for pollen, distributed in the ARS Pollen Bank as Oregon 1A.
Soil samples were hard to make because of the abundance of chert, sandstone, or quartz at almost all sites. Every soil sample was quite acidic with pH ranging between 3.96 and 5.80. The samples were low in phosphorous (3 to 20 ppm), low to moderate in potassium (32 to 123 ppm), and wildly variable for organic matter (3.1 to 56.8%). Azaleas growing on soils low in organic matter generally had roots scattered among the rocks or in cracks, while the high organic matter samples came from small pockets above and around rocks where leaves and other organic matter could accumulate and decompose.
In mid-October, we returned to collect seeds. We collected seeds from 10 of the sites we visited earlier and 2 additional sites in the Mark Twain National Forest in Missouri. Pods were ripening but few had already split open.

Rhododendron prinophyllum in Hawn State 
Park
Rhododendron prinophyllum growing on sandstone
bluffs under shortleaf pine in Hawn State Park,
Sainte Genevieve County, Missouri.
Photo by Sharon K. Dragula

Some of Missouri's more easily observed and attractive azalea populations occur in Hawn State Park, in Sainte Genevieve County, on bluffs just across Pickle Creek south of the campground (see photo), in Pickle Spring Natural Area, southwest of Hawn State Park in Sainte Genevieve County, and at the Spiva Azalea Park on State Highway 72 east of Fredericktown in Madison County.

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Deciduous Azaleas in Ohio
In May 1990, one of us (Larson) made two trips to southern Ohio to visit blooming populations of all three of Ohio's native azalea species. Rhododendron prinophyllum and its near likeness, R. periclymenoides , were in flower on the 8th of May and R. calendulaceum was blooming on the 18th. Rhododendron prinophyllum was found in both Scioto and Pike counties, R. periclymenoides only in Scioto County and R. calendulaceum in Pike County.
Together we visited another population of R. calendulaceum in June 1991. It is Ohio's northernmost population, located on private land near Sugar Grove, Fairfield County, and also one of the state's largest populations. We were too late to observe flowering, but the owner kindly showed us the population, in addition to photos of both yellow and orange blossoms. Seeds were collected from this site in late Sept. 1991.
Rhododendron prinophyllum and R. periclymenoides both had light pink flowers and the Pike County populations of R. calendulaceum included a range of yellow to orange flowers (see photo). Most plants were growing on north to northwest-facing slopes, often in soils with a pronounced surface layer of sand. Ohio azaleas generally grow on somewhat less rocky soils than do those from Illinois and Missouri; but, with two exceptions, Ohio soil test results were similar to those for sites farther west.

Rhododendron calendulaceum from 
Pike County, Ohio
A yellow form of Rhododendron calendulaceum
from Pike County, Ohio.
Photo by Richard A. Larson

At all but two sites in Scioto County, azaleas were growing on well-drained, acidic soils (pH 4.33 to 5.69), very low in phosphorous (1 to 5 ppm), low to moderate in potassium (44 to 64 ppm), and low to moderately high in organic matter (1.8 to 8.6%). It is important to note, however, that R. prinophyllum was growing at two locations in Scioto County on atypical soils with no signs of chlorosis. At these sites, the soils were neutral (pH 6.86 to 7.13), very low in phosphorous (2 to 3 ppm), moderate in potassium (57 to 72 ppm), and moderate to moderately high in organic matter (3.0 to 8.5%). It may be that limestone fill from nearby roads has modified these soils, but such influences were not obvious. These two populations were made available through both the ARS Pollen Bank and Seed Exchange (as Nile 1 and 1 Union) and we would be very interested to learn if these azaleas are especially well adapted to higher pH soils.

Rhododendron maximum in Ohio
In 1991, we turned our attention to the rosebay rhododendron, R. maximum . This species is one of the important parents of our hardiest rhododendrons and Schroeder (1985) discussed its potential to improve the adaptation of hybrids to extreme conditions. We all travelled to southern Ohio in mid-June and one of us (Larson) made an additional trip there and one to Jefferson County, in the eastern part of the state, later in June and another to collect seeds in late September. Altogether, we observed R. maximum in eight different areas, made pollen collections from six populations, and collected seeds from four. Rhododendron maximum , unlike the azaleas, grows on fairly moist sites with hemlock ( Tsuga canadensis [L.] Carr.), sweet birch ( Betula lenta L.), sugar maple ( Acer saccharum Marsh.), and beech ( Fagus grandifolia Ehrh.). In most populations, a large majority of plants did not flower either because of their small size or excess shade. For nearly all populations, flowering characteristics were quite uniform. Throughout the Ohio populations, peak flowering occurred during the last two weeks of June. Petal colors included only shades of light pink (see photo)(RHS Color Classes 56 and 73), fading to nearly white, except for the Jackson County population (distributed as Byer) which showed considerable variability for flower color including a few plants with a more intense pink color (petals RHS 74D, bud 72C). Most flowers also had a yellow-green blotch (RHS 150A). Floral diameter ranged from 3.0 to 5.0 cm.
Plants were growing on a mix of sand and organic matter, usually over sandstone or along sandstone cliffs. The soils were very acidic (pH 3.55 to 4.84), very low in phosphorous (1 to 5 ppm), except for the Byer population which had a reading of 104 ppm, low to moderate in potassium (28 to 94 ppm), and moderate to high in organic matter (4.5 to 16.4%). We suspect that excess organic matter is needed for the establishment of seedlings of this plant on sandy soils.

Rhododendron maximum in 
Fairfield County, Ohio
Rhododendron maximum at the Wahkeena
State Memorial, Fairfield County, Ohio.
Photo by Sharon K. Dragula

In Ohio, R. maximum often grows on beautiful, but nearly inaccessible sites, in ravines or otherwise difficult terrain. The most easily accessible population of this plant in Ohio can be observed along the road leading to the Wahkeena State Memorial in Fairfield County near Sugar Grove.

Conclusion
The azaleas and rhododendrons native to the midwestern states may have been overlooked in the past, but we believe they have potentially useful characteristics that will encourage their use in future breeding. Through the ARS Pollen Bank and Seed Exchange, we have made available a cross-section of species adapted to climatic extremes. We hope that these collections will be used to improve the stress tolerance of new hybrids and that they may also include new sources of adaptation to neutral soils. Please feel free to contact us concerning your experiences with these collections, or about other populations of native Rhododendron in the Midwest.
A complete list of collection sites and soil test results is available on request from the senior author.

Acknowledgements
There are many individuals and organizations that kindly helped us with this project. Our thanks go to all who helped, including Ray Smith and John Taylor of the US Forest Service in Illinois, Jody Eberly, Gary Houf, B. Erik Morse and James Murrell of the US Forest Service in Missouri, James H. Wilson of the Missouri Department of Conservation, Joe Manick and Paul W. Nelson of the Missouri Department of Natural Resources, Douglas Ladd of the Missouri Field Office of The Nature Conservancy, Allison Cusick, Charlie Foster and Rick Queen of the Ohio Department of Natural Resources, Paul Carringer of Ohio Camp Fire Girls, and Bill Pearson for identifying sites and arranging permission, to the curators of the herbaria at Iowa State University, Southern Illinois University, Southwest Missouri State University, and the University of Missouri for providing collection information, and the Research Foundation of the American Rhododendron Society for financial support.

Literature Cited
1.  Boge, D. The Ross-Boge Rhododendron macrophyllum expedition. J. Amer. Rhod. Soc. 40: 62-65; 1986.
2.  Boge, D. The Ross-Boge Rhododendron macrophyllum expedition, 1986. J. Amer. Rhod. Soc. 41: 62-65, 106; 1987.
3.  Boge, D. The Ross-Boge Rhododendron macrophyllum expedition, 1987. J. Amer. Rhod. Soc. 42: 162-167; 1988.
4.  McCullough, M.A. Exploring for the western azalea in southern California. J. Amer. Rhod. Soc. 39: 70-74;1985.
5.  Schroeder, H.R. Hybridizing rhododendrons in the midwest climate. J. Amer. Rhod. Soc. 39: 24-26; 1985.
6.  Skinner, H.T. In search of native azaleas. Morris Arboretum Bulletin 6: 3-10, 15-22; 1955.
7.  Steyermark, J.A. Flora of Missouri. Ames, Iowa State University Press, Ixxxiii, 1728 pp; 1963.

1 2121 Burnett Avenue, Ames, IA 50010
2 The Dawes Arboretum, 7770 Jacksontown Road SE, Newark, OH 43055