Proc.Fla. State Hort. Soc. 92:286-289. 1979.

THE NATIVE GRAPE SPECIES OF FLORIDA1

David J. Rogers and John A. Mortensen There are differences in the number of genera (Small in Dept. of EPO Biology, Univ. of Colorado, cludes two genera) (17), and the number and names of Boulder, CO 80309 and, species and divisions within the genus . Only one IFAS, Agricultural Research Center, University of , species, V. rotundifolia Michx., appears in all of the classi P. O. Box 388, Leesburg, FL 32748 fications listed. Some of the differences are purely nomen- clatural ones, but others are more questions of interpreta Additionalindex words, classification, Vitis, . tion on what constitutes a species. The classical difference between "splitter" and "lumper" in classification is exhibited Abstract. Because genetic resources of wild grapes fre by Bailey (2) and Duncan (6). Small (17) was probably quently enter into the germplasm of some of our cultivated the most knowledgeable about the variability and ecology grapes, it is important to produce a classification that recog of the species because his field work was carried out over a nizes the relationship and variations of the native wild number of years, and he made more personal observations species alongwith attributes that might be useful in breed over the growing seasons than any of the other workers ing. We recognize two muscadine and three bunch grape in Table 1. But all of them left numbers of uncertainties species (one with five subspecies) in our classification. Eco and confusion about the classification of the wild species of logical differences are significant not only for classification, Vitis. It is the purpose of this paper to reclassify the species but also to aid in selection of environmentally adapted wild to bring the work as up to date as possible while recogniz material in breeding programs for different areas ofthe ing that many more observations must be made before the State of Florida. final decision on the "correct" classification can be made. Rogers and Mortensen worked together at Leesburg from June to November, 1978 while the former was on sabbatical. The native grapes of Florida have provided valuable germplasm for improvement of cultivated species in all of the southeastern United States. A selection of Vitis mun Materials and Methods soniana Simpson ex Munson provided the genes for perfect flowers (5) in muscadine grape breeding. Vitis smalliana Herbarium specimens, photographs, and preserved fruit Bailey (F. aestivalis subspecies smalliana in this classifica ing materials were made of the wild species grown in the tion) was used as a species background for the cultivars vineyard at the Agricultural Research Center, IFAS, Lees 'Blue Lake' and 'Norris', conferring resistance to Pierce's burg. These specimens were used to compare with wild- disease. V. shuttleworthii House was used in the cultivar growing members of the same species. The herbarium 'Stover'. Others have been used in various programs of collection of any included parts of the youngest breeding with varying degrees of success. Because of these shoots, canes, and mature older stems with leaves to demon important contributions to Florida's growing interest in a strate variability within and between ; if mature grape industry it was necessary to review the taxonomic fruits were available, FAA-preserved specimens were made. status of the native species to determine whether or not the Seeds were extracted and later germinated to provide extant classification is reflective of the existing genetic re seedling specimens as well as greenhouse-grown plants for sources. By comparing various classifications that include comparison. Standardized field data sets of each collection the Florida species, one may see that there is disagreement were gathered at the time of collecting, to include: (A) between various authorities and that some resolution of color of canes and shoots, pubescence color on canes and the differences needs to be made. Table 1 summarizes the shoots, tendril color both young and mature; (B) leaf species accepted by the most prominent students of grape characters such as upper surface texture, surface direction classification for those geographic regions including Florida. (flat, V-shaped, or cupped downward); (C) berry and cluster shapes, color and size of mature berries, presence

iFlorida Agricultural Experiment Stations Journal Series No. 2106. or absence of bloom, flavor, pulp consistency; (D) habitat Supported by National Science Foundation Grant DEB 77-15070 to description, estimates of frequency and cover. the first author. Collections were made by the first author in all sections

Table 1. Number of Florida species of Vitis recognized by various authorities.

Planchon, 1887 Munson, 1909 Small, 1933 Bailey, 1934 Duncan, 1967

Euvitis Euvitis Vitis Euvitis Vitis V. aestivalis V. candicans V. aestivalis V. cordifolia V. aestivalis V. cordifolia V. cinerea, var. V. cinerea V. illex V. shuttleworthii V. coriacea floridana V. cordifolia V. rufotomentosa V. rotundifolia Muscadinia V. cordifolia V. coriacea V. shuttleworthii V. vulpina V. rotundifolia V. coriacea V. rufotomentosa V. simpsonii V. simpsonii V. simpsonii V. smalliana Muscadinia Muscadinia V. sola V. rotundifolia M. rotundifolia Muscadinia V. munsoniana M. munsoniana V. rotundifolia V. munsoniana

one genus one genus two genera one genus one genus two subgenera two subgenera no subgenera two subgenera no subgenera 3 spp. bunch 5 spp. bunch 6 spp. bunch 7 spp. bunch 3 spp. bunch 1 sp. muscadine 2 spp. muscadines 2 spp. muscadines 2 spp. muscadines 1 sp. muscadine

286 Proc. Fla. State Hort. Soc. 92: 1979. of the state, guided by land form type and temperature Suwanee River to the St. Marys River). isotherms (Fig. 1) (3), vegetation maps (4) and soil maps 1. V* rotundifolia (8). Particular effort was made to collect in the type locali ties for species first described from Florida. 2. Berries smaller, generally less than 1 cm, with very little pulp. Clusters larger, usually 12 to 30 berries. Key to the Subgenera, Species and Subspecies Exocarp thinner and dark purple to black, without lenticels. Seeds smaller, 6 mm or less. Found almost 1. Bark of mature stems smooth, unbroken, glabrous, with exclusively in peninsular Florida, south of the Su- frequent lenticels, particularly at the nodes. Pith con wanee-St. Marys River line. tinuous, without diaphragm at the nodes. Tendrils simple, unbranched. Berries in loose, few-membered 2. V. munsoniana clusters, each berry easily separating from its pedicel. Bark of mature stems shredding in longitudinal strips (Subgenus Muscadinia). or squares, with diaphragm at the nodes. Tendrils fre 2. Berries large, pulpy, usually 1.5 cm or greater. quently branched. Berries in relatively large, many- Clusters small, usually 2 to 8 berries. Exocarp either membered clusters. (Subgenus Vitis) bronze or purple, thick and tough, frequently lenti- 3. Leaves (even the young ones) glabrous or essentially cellate. Seeds 8 mm long or longer. Species found so; cordate, rarely 3- or 5-lobed. Found principally almost exclusively in the north and western parts in north and west Florida. of Florida (generally, N. and W. of a line from the 3. V. vulpina

25

Fig. 1. Major landforms of Florida and isotherms for number of hours per year below 24°F. Adapted from Brooks and Rappenecker (3). Proc. Fla. State Hort. Soc. 92: 1979. 287 Leaves generally pubescent, frequently heavily so; in sinuses and "fig-leaf shape; outline, either shield-shaped, ovate, obovate or reni- margins entire or obscurely and form, from slightly longer than broad to broader irregularly toothed. Mostly on than long. deep sandy soils. 5d. F. aestivalis ssp. smalliana 4. Leaves thickened, with whitish (infrequently light brown), heavy, felted tomentum beneath, 8. Leaves with pointed lobe tips, 3-, the upper surface rugose, the leaf with a down 5-lobed; margins serrate. On ward curve from the midrib. Fruit large, 1 cm or loamy sand soils. more. Distributed in the peninsula alone, its 5e. V. aestivalis ssp. simpsoni northern limits on a line from about Levy County (Cedar Key) on the west to Volusia Descriptions of Species and Subspecies County on the east, most frequently in wet, low- lying regions. 4. V. shuttleworthii 1. Vitis rotundifolia Michaux (12). "Muscadine".

4. Leaves with a thin, flattened blade, with rusty This species, with roundish, glossy green, very deeply tomentum beneath (but never felted), the upper serrate leaves, is the most widely cultivated species of grapes surface smooth. Berries smaller, usually less than in the southeast. Its large berries fall easily from the 1 cm diam. Distributed throughout the state cluster, one of the greatest problems for its use as a com and north to at least New Jersey, in mesic condi mercial fruit. The cultivar 'Scuppernong' was selected very tions. Five subspecies. 5. F. aestivalis early after European contact in North America, and is supposed to be named after a river in North Carolina. F. 5. Leaves generally orbicular, seldom indented rotundifolia has been widely used in the breeding of all or lobed; pubescence mostly brownish. Con the present-day muscadine cultivars. Crosses with any of fined to mostly northeastern counties of the species of subgenus Vitis are difficult and most sterile. Nassau, Duval, Clay, and St. Johns. However, there have been recent successes (7, 14) that make 5a. F. aestivalis ssp. sola breeding a much more likely possibility to improve the culti 5. Leaves at least shouldered, most frequently vars. One of its greatest values is that the species in general at least 3-, and often 5-lobed; pubescence is resistant to Pierce's disease, Phylloxera, and nematodes. brown, tawny or rusty. Distribution over much of state. 2. Vitis munsoniana Simpson ex Munson (16). "Bullace".

6. Leaves most commonly shield-shaped, 3-, Although this species was recognized nearly one hundred 5-lobed, the tips of lobes pointing for years ago, it has not been considered separate from V. ro ward; margins very coarsely serrate; tundifolia by a number of workers. Duncan (6) did not leaves of young, vigorous canes fre recognize V. munsoniana in his study of southeastern vines. quently very deeply 5-lobed and much One reason is that in their vegetative state, the vines of the larger than the average on the plant. two are nearly indistinguishable, and herbarium specimens Distribution extending from Georgia are of little value for this distinction, unless mature fruits and Alabama into northern tiers of are collected with the specimen. However, V. munsoniana counties and down east coast at least to has an almost completely distinct geographic range in the Volusia County. peninsula of Florida, whereas V. rotundifolia is found to the 5b. V. aestivalis ssp. aestivalis north and west of the bullace grape's range. Moreover, the ripening period of V. munsoniana extends from August to 6. Plants with nearly the same types of leaf December, whereas F. rotundifolia ripens only in August variations, but margins either serrate, and September. Budbreak occurs in V. munsoniana a full serrulate, or smooth. Distribution mostly month before it does in V. rotundifolia, but defoliation in in central and southern part of the state. the fall is later than V. rotundifolia. 7. Leaves longer than broad, with There is a small-leaved form of F. munsoniana that sharply pointed lobes, the central occurs infrequently throughout the range of the species. lobe elongate, the 2 uppermost This form, with pronouncedly smaller leaves, but other lateral lobes frequently pointed out wise similar to the majority of plants, has been given a wards. Inflorescence large, lax, with name that was never published, therefore, will require de apparently angled secondary scription and publication for validation. The form was branches. Seeds very small, 4.8 mm called Muscadinia pygmaea by a collector on the label of an long or less. Distribution in coastal herbarium specimen (J. B. McFarlin #5707) in the United lowlands from Manatee County on States National Herbarium. We believe that this form west coast south to Dade County and should be recognized and will, after further investigation, up east coast to Brevard County. probably designate it as F. munsoniana forma pygmaea. 5c. V. aestivalis ssp. divergens Most of the self-fertile cultivars of muscadines grown 7. Leaves as broad as long, either with today can be traced back to at least some F. munsoniana sharply pointed or rounded lobe tips, in the parentage. but if sharply pointed, the points facing forward. Seeds 5 mm long or 3. F. vulpina L. (11) longer. Plants largely of the central highlands, on either deep sand or This species name has been unfortunately misinter loamy sand. -, preted by a number of taxonomists and viticulturists over the years. Rogers recently examined the type specimen of 8. Leaves with uniformly rounded F. vulpina (at the Linnean Herbarium in London) and lobe tips, most often with deep the types of F. cordifolia in the Phanerogamic Herbarium

288 Proc. Fla. State Hort. Soc. 92: 1979. of the Natural History Museum in Paris with the result Vitis aestivalis Michaux subspecies aestivalis. that, although the name V. cordifolia has been in wide sola (Bailey) spread use for some time, it must be made a synonym of simpsoni (Munson) V. vulpina according to the International Rules of Botanical smalliana (Bailey) Nomenclature. Bailey's Vitis illex, erected as a new status divergent (Rogers)2 for Vitis cordifolia var. sempervirens Munson (2), is simply a minor variant of V. vulpina and is, therefore, treated as a The formal transfer of these names with citation of the synonym of V. vulpina. A full scale discussion of the intri types, etc., will be done in the generic monograph, which is now in preparation. We have placed V. rufotomentosa cate nomenclature problems involved is not appropriate to Small in synonymy under V. aestivalis ssp. simpsoni. this paper, but will be given in a monographic treatment Some of the subspecies, such as simpsoni, have been used of the North American species of grapes. extensively in breeding programs to bring resistance or Vitis vulpina is a clear-cut, wide-ranging eastern species tolerance to Pierce's disease. For example, simpsoni was with very little intraspecific variation. It has been used in used to develop 'Lake Emerald', 'Norris', and newer selec France in breeding for Phylloxera-resistant rootstocks, but is tions of improved quality with potential as scion or root- not now in use. The very small berries and undesirable stock cultivars. Subspecies smalliana was used in the develop flavor are hardly useful traits, but it might, however, have ment of cultivars 'Blue Lake' and 'Norris'. Some hybridiza some resistance to Pierce's disease and Phylloxera (13) tion of V. aestivalis is probably in the background of that could be useful in breeding rootstocks. Munson's V. bourquiniana varieties such as 'Herbemont' and 'Lenoir'. The other two subspecies have not, to our 4. Vitis shuttleworthii House (9); see Bailey (2). knowledge, been used in breeding programs, but we think The nomenclature of this species is quite involved, and that they soon will be, particularly since we have indicated is well explained by Bailey, q.v. The relation of this species their relation to the parental species, which has been used to Vitis candicans Engelm., a Texas species, was recognized successfully. These two, subspecies sola and divergens, each early by Bailey (1) and by Koch even earlier (10). However, have qualities that should be useful in development of its great geographic disjunction from V. candicans, and bunch grapes for Florida vineyards. slight variations in morphology, physiology, and phenology (it blooms and fruits earlier and is more susceptible to winter Literature Cited cold and alkaline soil than V. candicans) are sufficient to iThis is a provisional name for a new subspecies of V. aestivalis. warrant designation as a separate species. Also, the ripe 2. . 1934. Vites peculiares ad Americam Borealem. Gentes fruit of V. candicans has a fiery pungency, but the flavor of Herbarum III (Fasc. IV): 151-244. V. shuttleworthii is free of this. Although the fruit is some 3. Brooks, H. K. and C. Rappenecker. 1964. in Raisz, E. & Associates, Atlas of Florida, with text by Dunkle, J. R. Univ. of Fla. Press, what insipid, it has been frequently used in breeding with 52 pp. other bunch grapes because of its disease and insect re 4. Davis, J. H. 1943. The Natural Features of Southern Florida. Geol. sistance, vigor, and size of berry. It was used in the breed Bull. 25, State of Florida Dept. of Conservation. ing of 'Stover' bunch grape, the most promising cultivar in 5. Dealing, C. 1917. Muscadine grape breeding. J. Hered. 8:409-424. Florida. The species has also been useful in developing root- 6. Duncan, W. H. 1967. Woody vines of the southeastern states, Vitacae. Sida 3:22-32. stock cultivars with resistance to Pierce's disease and nema- 7. Dunstan, R. T. 1962. Some fertile hybrids of bunch and muscadine todes. grapes. /. Hered. 53:299-303. 8. Henderson, J. R. 1939. The Soils of Florida. Univ. of Fla. Agric. Expt. Sta. Bull. No. 334. 5. Vitis aestivalis Michaux (12). 9. House, H. D. 1921. Am. Midi. Nat. 7:129. 10. Koch, K. H. E. 1869. Dendrologie, Vol. 2:550. Bailey (2) established section Aestivales in the genus 11. Linnaeus, C. 1753. Species Plantarum. for all the species he recognized as having affinity with V. 12. Michaux, A. 1803. Flora Boreali-Americana, Vol. 2:230, 231. aestivalis, and separated many of the geographic variants 13. Munson, T. V. 1909. Foundations of American Grape Culture. as species. In Florida, he named five separate species in Orange Judd Co., New York. this section, but excluded V. aestivalis itself, stating that 14. Patel, G. I. and H. P. Olmo. 1955. Cytogenetics of Vitis: I. The hybrid V. vinifera x V. rotundifolia. Am. J. Bot. 42:141-159. V. aestivalis did not extend south of Georgia. After con 15. Planchon, J. E. 1887. Monographic des Ampelidees Vrais. in D.C siderable field work and studies of some of the variants in Monographia Phanerogamarum V:305-368. the vineyard, we believe that the more realistic classification 16. Simpson ex Munson. 1887. in Proc. Soc. Prom. Agr. Sci. 59. reestablishes V. aestivalis in Florida, designating the geo^ 17. Small, J. K. 1933. Manual of the Southeastern Flora. Publ. by the Author, N.Y. pp. 835-838. graphic variants as subspecies. We recognize, therefore, the following:

Proc. Fla. State Hort. Soc. 92: 1979. 289