FEATURE The Muscadine : Botany, , History, and Current Industry William C. Olien1 U.S. Department of Agriculture-Agricultural Research Service, Small Fruit Research Station, Box 287, Poplarville, MS 39470

The muscadine grape ( rotundifolia grape found in India has similar characteristics is the name of a specific . Michx.) was the first American grape spe- to the Muscudinia (Syamal and Patel, 1953) The natural range of V. rotundifolia ex- cies to be cultivated. This fruit has a long and may reflect the pre-ice age distribution of tends from Delaware to central and history in commercial and backyard culture. this group. along the Gulf of Mexico to eastern Picking muscadines to enjoy as juice along Muscadinia have 40 somatic chro- (L.H. Bailey Hortorium, 1976; Dearing, with muscadine pie on grandmother’s back mosomes (2x = 2n = 40) and are charac- 1938; Munson, 1909; Weaver, 1976). The porch has a strong place in childhood mem- terized by fruit borne in many clusters with species extends north along the ories of many native Southerners. The oldest few berries per cluster, formation of an ab- River to Missouri and near the Appalachian and most consistent commercial interest in scission zone between the fruit and rachis, Mountains from the east and west. Temper- muscadines has been in , but juice and smooth thin bark that is adherent on young atures in this region seldom go lower than fresh fruit markets have also been devel- wood and separates in scales from older wood, - 12C and more rarely to - 18C (U.S. Dept. oped. Muscadines so differ from “bunch” unbranched tendrils, dense wood, and con- of Agriculture, 1973). These vines do best grapes genetically, anatomically,, physiolog- tinuous pith (L.H. Bailey Hortorium, 1976; on fertile sandy loams and alluvial soils, and ically, and in taste that they should be con- Einset and Pratt, 1975; Hedrick, 1908; Mun- grow poorly on wet and heavy soils. Natural sidered a separate fruit. son, 1909; Williams, 1923). In contrast, Eu- populations are found in shady, well-drained The muscadine industry is currently ex- vitis grapes have 38 somatic chromosomes bottom lands along rivers that are subject to panding throughout the southeastern U.S. as (2x = 2n = 38), branched tendrils, many neither extended drought nor waterlogging interest in grapes and wine increases nation- berries per fruit cluster, no abscission zone (Hedrick, 1908; Munson, 1909). ally. Commercial muscadine range between the berry and rachis, striated bark Wild V. rotundifolia vines are functionally in size from < 1 ha to hundreds of hectares, on young wood, thick rough bark that peels - dioecious (polygamous) due to incomplete sta- and may be a primary or secondary source in strips on old wood, less-dense wood than men formation in female vines and incomplete of income to the grower. There is particular Muscudinia, and pith interrupted by dia- pistil formation in male vines (Dearing, 1948; interest in this fruit for small and part-time phragms at nodes. Hedrick, 1908). Male vines account for 60% farm operations and as an alternative crop Muscudinia is a much smaller group than to 75% of the wild muscadine population for agronomic growers. An important recent Euvitis, and is comprised of three known (Dearing, 1938; Husmann and Dearing, 1916). development has been the establishment of species. Vitis munsoniana Simpson ex Mun- Female vines bear fruit in numerous clusters a major muscadine juice and processing son (common names Bird Grape, Everbear- of from one to 40, but more commonly four in Mississippi with a satellite plant in North ing Grape, Mustang Grape, Little Muscadine to 10, thick-skinned berries containing two to Carolina. Growth of the muscadine industry Grape) and V. popenoei Fennell (common six large (Husmann and Dearing, 1916; is supported by research to improve name, Mexican Muscadine Grape) (Munson, Young, 1920). The vines are late in breaking management, processing methods, and cul- 1908; U.S. Dept. of Agriculture, 1973; bud in the spring and require a long season, tivars through breeding. Weaver, 1976; Winkler et al., 1974) are not generally > 100 days, to mature the fruit (Hed- important commercially, but are possible rick, 1908). BOTANICAL BACKGROUND sources of genetic variation for breeding pro- Most authorities divide Vitis into the subge- grams. Vitis munsoniana is likely a semi- EARLY MUSCADINE nera Euvitis Planch. (the familiar European and tropical variant of V. rotundifolia, native to Muscadines have long been harvested from American bunch grapes that include V. vini- Florida and a narrow coastal band along the wild and semicultivated vines (Hedrick, fera L. and V. labrusca L.) and Muscadinia Gulf of Mexico from Florida to Texas (Hus- 1908). According to Hedrick, Native Amer- Planch. (sometimes referred to as berry grapes) mann and Dearing, 1916). Vitis munsoniana icans used the fruit, and the earliest Euro- (Dearing, 1938; Winkler et al., 1974). How- bears clusters of eight to 30 relatively small peans to reach America noted the abundance ever, there has been a long-standing contro- berries with thin skin and small seeds, but of V. rotundifolia. Captain John Hopkins re- versy, with some authors placing Muscadinia poor fruit quality (Dearing, 1947). Vitis po- ported that Spanish missionaries in Florida as a separate genus (Bouquet, 1980; Olmo, penoei is a tropical species native to southern were making muscadine wine in 1565. He 1986; Small, 1913). Fossil evidence suggests Mexico (Fennell, 1940). also noted that Amadas and Barlowe re- that Muscadinia was widely distributed over The third species, V. rotundifolia Michx. ported the abundance of these grapes in coastal the North American, European, and Asian (common names: Muscadine, Bullace, Bull on their first voyage to this continents before the last ice age (Bouquet, Grape, Bullet Grape, Southern Fox Grape), continent in 1584. The latter described the 1980). Thus, Muscadinia may be an ancestral is the only commercial Muscadinia grape, land “so full of grapes . . . on the sand and genus that led to vitis, adapted to temperate and the name “muscadine” is reserved ex- on the green soil, on the hills as on the plains, climates,. and Ampelocissus, adapted to tropi- clusively for this species. The majority of V. as well as on every little shrub as also climb- cal climates, as the ice age receded. A wild rotundifolia vines in the wild bear dark fruit. ing towards the tops of tall cedars, that I A very old name for these grapes is “Bul- think in all the world the like abundance is lace” or “Bullis”, or less commonly “Bull” not to be found”. Received for publication 19 May 1989. The cost or “Bullet” grapes (Gohdes, 1983; Hedrick, Muscadines have been cultivated in vine- of publishing this paper was defrayed in part by the payment of page charges. Under postal regu- 1908). Light-colored bronze-fruited geno- yards since the mid-18th century (Reimer, lations, this paper therefore must be hereby marked types are occasionally found in the wild and 1909). The first cultivars were simply selec- advertisement tsolely to indicate this fact: are often referred to generically as “scup- tions propagated from the wild (Reimer, 1Present address: Dept. of Horticulture, Clemson pernongs”. However, there are many culti- 1909). Although there were 35 to 40 named Univ., Clemson, SC 29634-0375. vars of bronze muscadines, and ‘Scuppernong’ muscadine cultivars in 1920, only a few were

732 HORTSCIENCE, VOL. 25(7), JULY 1990 grown extensively (Young, 1920). Better- A second USDA-ARS muscadine breeding rootstocks for Euvitis grapes (Davidis and Olmo, known early cultivars and approximate dates program was conducted at Meridian, Miss., 1964; Husmann and Dearing, 1916; Winkler introduced were ‘Hopkins’ (1845), ‘Flow- from 1941-1965 by N.H. Loomis. Mus- et al., 1974). In contrast to Euvitis, musca- ers’ (1800), ‘James’ (1866), ‘Memory’ cadine breeding was initiated by the Geor- dines root poorly from woody cuttings (Goode (1868), ‘Mish’ (or ‘Meisch’) (1846), ‘Scup- gia Experiment Station in 1909 (Stucky, et al., 1982; Husmann and Dearing, 1916). pernong’ (mid-1700s), and ‘Thomas’ (1845) 1919). In addition, various private individ- Commercial practice is to propagate either by (Husmann and Dearing, 1916; Reimer, 1909). uals have conducted muscadine breeding layering (Woodroof, 1936) or by mist-propa- All early cultivars were female types and re- efforts, most notably the Euvitis x V. ro- gation of softwood cuttings (Goode and Lane, quired pollinator vines for optimum fruit set tundifolia hybridization efforts of T.V. 1983). 1H-indole-3-butyric acid (IBA) in- (Young, 1920). Munson (1909) in Texas, and Olmo (1986) creased root system quality of cuttings taken The cultivar eventually named ‘Scupper- and coworkers in California. Active mus- prior to July, but did not increase root initia- nong’ was the first native American grape to cadine breeding programs for fruiting cul- tion or shoot growth (Goode and Lane, 1983). be cultivated (Hedrick, 1908; Reimer, 1909). tivars are currently being conducted in Vines are sold bare-root or in “1-gallon” (3.8- ‘Scuppernong’ was the dominant cultivar Florida (Bates et al., 1980; Mortensen, liter) nursery pots. Micropropagation methods ‘grown from the mid-18th century until as 1971), (Lane, 1978), and North are currently being developed by several groups recently as 1947, and it remains the most Carolina (Goldy, 1988; Goldy et al., 1988). (Gray and Fisher, 1985; Griffin and Graves, widely known muscadine (Reimer, 1909; Progress in breeding up to the early 1970s 1989; Lee and Wetzstein, 1989; Sudarsono Dearing, 1947; Woodroof, 1934). Popular was summarized by Einset and Pratt (1975) and Goldy, 1988). culture often gives credit for discovery of the and Schwartz (1976). ‘Hunt’ was intro- Planting and establishment. General rec- original ‘Scuppernong’ vine to Sir Walter duced in 1920 by the Georgia Experiment ommendations on site selection, soil prepara- Raleigh’s colony when they landed on Roa- Station (Stucky, 1919) and had improved tion, planting methods, and vineyard noke Island, N.C. (Gohdes, 1982; Hedrick, yield and fruit quality, but was female and management are given in several muscadine 1908). However, Reimer (1909) concluded required companion pollinators. The de- production manuals (Hegwood et al., 1983; that the original ‘Scuppemong’ vine was found velopment of perfect-flowered, self-fertile Ferree et al., 1983; Poling et al., 1987; U.S. by Isaac Alexander in the mid-18th century cultivars was a major goal of early breed- Dept. of Agriculture, 1973). While mature along the Scuppernong River in Tyrrell ing programs. The first self-fertile geno- muscadine vines are noted for vigorous growth, County, N.C. The vine was soon widely types were reported by the North Carolina- they can be difficult to establish after planting propagated and was likely carried to Roa- USDA breeding program in 1917 and the (Husmann and Dearing, 1916). Recent studies noke Island some time in the late 18th cen- first cultivars were released in 1948 (Dear- have determined that size, shape, and wall tury. This cultivar was initially known by ing, 1917, 1948). roughness of the planting hole, and severity of several names, especially the “Big White A long-standing goal of both Euvitis and summer pruning in first-season vine training, Grape”, and it was not until 1811 that the V. rotundifolia breeding programs has been have a marked effect on establishment and name ‘Scuppernong’ was assigned by Calvin development of hybrids between these growth of muscadines (Olien, 1989). Jones, an editor for The Star newspaper of groups, combining fruit quality from V. Training and trellis. Traditionally, musca- Raleigh, N.C. The name was chosen be- vinifera with disease resistance and envi- dines were grown on extensive overhead arbor cause of the numerous plantings of this grape ronmental adaptation of muscadines (Bou- systems, often without pruning or manage- along the Scuppernong River and around Lake quet, 1980; Einset and Pratt, 1975; Goldy ment (Husmann and Dearing, 1916; Newman, Scuppernong. The word scuppernong is ap- et al., 1988; Lane, 1978; Munson, 1909; 1907). Arbor height was typically 2.1 m, with parently a corruption of “ascuponung”, an Olmo, 1986). Hybridization is difficult due eight arms radiating out from single trunks, Algonquin Indian word for “place of the to differences in chromosome number, but spaced at 4.6 x 4.6 m. Newman (1907) rec- Sweet Bay plant” [Magnolia virginiana L. has been successful when the muscadine ommended a four-wire vertical trellis (4-WVT) (M. glauca L.)], found abundantly along the was used as the male parent (Davidis and over the arbor system for commercial produc- Scuppernong River. Olmo, 1964). Most hybrids have been ster- tion because it was easier to manage and har- ‘Scuppernong’ was especially valued for ile, but a few have a low level of fertility. vest. However, the vigorous growth habit of its hardiness, tolerance of neglect, and qual- In addition to standard breeding tech- muscadines soon led to recommendations of ity as a table and wine grape. Bronze mus- niques, tissue culture and protoplast fusion two or three wires in vertical systems to reduce cadines were soon recognized as superior for methods are being employed with the hope shading of lower cordons and consequent in- wine because the nonacylated diglucoside of developing fruitful hybrids through crease in growth, yield, and fruit quality pigments of dark muscadine backcrossing programs to develop both V. (Dearing, 1938; Husmann and Dearing, 1916; grapes are unstable and easily oxidized, in vinifera and V. rotundifolia cultivar types Savage, 1941; Young, 1920). Generally rec- contrast to the monoglucoside and acylated (Goldy et al., 1988; D. Gray, personal ommended spacing was 6.1 to 6.7 m in-row monoglucoside anthocyanin pigments of Eu- communication; Lee and Wetzstein, 1988). and 3.7 to 4.6 m between rows. In a 20-year vitis grapes (Ballinger et al., 1974). Re- No Euvitis x Muscadinia hybrids have been study, Brightwell and Austin (1975a) found cently, wild muscadine genotypes have been released as fruiting cultivars, but two hy- that long-term yields of ‘Hunt’ were lowest on identified with mono- and diglucoside forms brids were recently released as disease-re- single-wire trellis, intermediate on a 2-WVT, of anthocyanin pigments (Goldy et al., 1989). sistant rootstocks for Euvitis grapes (Lider and highest on the arbor system, if managed Continuous vegetative propagation of this et al., 1988a, 1988b; Walker et al., 1989). with adequate pruning. However, highest yield cultivar for nearly 250 years, and probable The development of a seedless muscadine in the first two bearing years was obtained introduction of other bronze muscadines under for table use is a major goal in current breeding with the 2-WVT system, an important advan- the same name, has resulted in the formation programs. Embryo rescue from standard sex- tage in generating income to offset establish- of a number of strains of ‘Scuppernong’ ual crosses of seedless Euvitis grapes (female ment costs. In-row vine spacing from 6.1 to (Woodroof,. 1934). ‘Scuppernong’ vines in parent) and V. rotundifolia (pollen parent) 6.7 m had no effect on yield/area, but be- production have been reported as old as 150 (Goldy et al., 1988), and protoplast fusion tween-row spacing of 3.0 m gave higher long- years (U.S. Dept. of Agriculture, 1973). methods (D. Gray, personal communication; term yields/area than either 3.7 or 4.3 m be- Lee and Wetzstein, 1988) are being used in tween rows (Austin and Bondari, 1989; BREEDING PROGRAMS an effort to develop seedless hybrids with V. Brightwell and Austin, 1975b). When a two- Breeding programs to obtain improved rotundifolia character. wire horizontal trellis, based on Geneva Dou- cultivars were initiated at Willard, N.C., ble Curtain (GDC), was compared with 2- around 1907 by Dearing (1917) and Detjen MUSCADINE CULTURE AND WVT, the GDC system averaged 11% greater (1917a) as a cooperative effort between the PHYSIOLOGY yields and 0.6°Brix higher soluble solids for USDA-ARS and North Carolina State Univ. Propagation. Muscadine grapes are not 31 cultivars over the first 3 to 5 crop years and continues today as a university project. grown on rootstocks, and are incompatible as (Andersen et al., 1985; Andrews, 1981). R.P.

HORTSCIENCE, VOL. 25(7), JULY 1990 733 Lane (personal communication) noted a 63% ‘Thomas’ were pruned to one-, two-, and Dearing, 1916). Thus, before the devel- yield increase for GDC over a single-wire trel- three-, or four-bud spurs or for mature ‘Hunt’ opment of perfect-flowered cultivars, 11% of lis, but pruning costs were 29.7 and 7.4 work- pruned to two- to four-bud spurs, with and the vineyard was occupied by nonbearing vines. ing hours/ha (12 and 3 hr/acre), respectively. without 50% spur thinning, or to seven- to Today, perfect-flowered cultivars are used as Muscadines grown on GDC are not main- eight-bud canes (Loomis et al., 1949). Lane pollinizers, greatly increasing yields per hec- tained as two separate “curtains”, but shoots (1977b) compared 10 cultivars over the first tare. from parallel cordons are allowed to grow to- five fruiting years pruned to three-bud spurs Muscadines pollen is dry and is probably gether. This arrangement creates a shaded, hu- or nine-bud canes, with equal number of buds transferred to pistillate cultivars almost en- mid canopy in the fruiting zone and in some retained in each treatment. Yield of vigorous tirely by insects (Dearing, 1938; Detjen, regions results in an unacceptable amount of cultivars was consistently greater with cane 1917b). Dearing (1938) reported that the most fruit rot (G.W. Krewer, personal communi- pruning. Other cultivars showed an occa- efficient pollinator is the small mining bee cation). Some growers have hedged the lateral sional yield increase with cane pruning, but (Halictus stultus Cress.), followed by the shoots to near the fruiting zone soon after ver- spur pruning was never superior in these green bee (Agapostemon splendens Lep.), aison to aid spray penetration and decrease young vines. Higher yields with cane prun- gray bee (Magachile sp.), and small bum- canopy humidity. There have been no research ing were a result of greater bearing surface. blebee (Bombus irnpatiens Cress.), and that evaluations of the possible benefits or hazards Results of this study also indicated the onset the honeybee (Apis mellifica L.) is not an (such as increasing risk of fall cold injury) of of alternate bearing in cane-pruned vines after efficient pollinator of muscadines. Beetles this practice. Leaf removal has become a stan- the fourth treatment year. Annual pruning have also been noted as possible muscadine dard practice to reduce canopy density of bunch must be accompanied by regular spur thin- pollinators, particularly Copidita thoracica grapes (Winkler et al., 1974), but this has not ning. Excessive spur development over sev- F. and the soldier beetle (Chauliognathus been tried in muscadines. eral years leads to shading, poor canopy marainatus F.) (Dearing, 1938; Detjen, 1917; In current commercial production, musca- structure, and poor fruit quality, similar to Armstrong, 1936). Dearing (1938) estimated dines are trained to single trunks with per- the problems with no pruning (Lane, 1977b; that fruit set of pistillate cultivars was nor- manent cordons on 1- or 2-WVT or on GDC. Loomis et al., 1949). mally 7% to 10%, but could be increased to The most common spacing is 3.7 m between Current practice is to prune to two- to four- 20% to 30% with proper pollination. The rows × 6.1 m in the row. Optimum trellis bud spurs, up to six-bud spurs for vigorous primary means of pollen transfer in perfect- design and training system depends on a bal- shoots, in combination with occasional light flowered muscadine cultivars is not known. ance of management costs, yield, and price spur thinning (Hegwood et al., 1983; Ferree Winkler et al. (1974) concluded that self- per tonne of fruit. The GDC system is rec- et al., 1983; Poling et al., 1987; U.S. Dept. pollination is the primary method of pollen ommended in Mississippi (C.P. Hegwood, of Agriculture, 1973). Mechanical hedging transfer in V. vinifera grapes, and that insect personal communication), while the single-wire with follow-up hand-pruning is practiced by and wind transfer of pollen were relatively trellis is most commonly recommended in growers with large operations. Mainland et unimportant. Various beetles and flies, but Florida, Georgia, and the Carolinas (R.G. al. (1982) reported that hedging a 20 × 20- not bees, have been noted at anthesis in mus- Goldy, R.P. Lane, and J.A. Mortensen, per- cm square around the cordon without addi- cadine test vineyards in North Carolina (R.G. sonal communication). tional hand-pruning resulted in increasingly Goldy, personal communication). Fruit set Management of fruiting wood. Dormant dense growth over 3 years that made in six perfect-flowered muscadine selections pruning was originally believed to be inju- difficult. However, hedging to 20 × 40-cm ranged from 11.3% to 23.3% (Goldy, 1988). rious, but several early authors disputed this and alternating the long axis between a hor- An initial set of berries develops opposite claim (Husmann and Dearing, 1916; New- izontal and vertical direction each year main- leaves at nodes 3 through 6 from the base of man, 1907; Reimer, 1909; Young, 1920). Ac- tained a manageable canopy without current season shoots, as in many grapes (Pratt, cording to Dearing (1938), lack of pruning decreasing yield. 1971). However, muscadines have a marked decreased fruit quality and increased alternate A further important task of dormant-pruning tendency to develop additional flower buds on bearing, early defoliation, and susceptibility to is removal of tendrils that encircle cordons or lateral shoots that can flower and set fruit in winter injury. Late winter or early spring prun- spurs. Muscadine tendrils become extremely the year they are initiated (Husmann and Dear- ing results in highly visible “bleeding” of sap tough and will girdle spurs and even the main ing, 1916; Young, 1920). This tendency re- from pruning wounds. Newman (1907) noted cordon if not removed each year. Yield in- sults in uneven ripening with fruit ranging from heavy bleeding in vines pruned in February or creases associated with pruning will be lost if a few millimeters in diameter to fully mature later and recommended pruning earlier than equal care is not given to removal of circling at harvest. Berries from the initial fruit set ripen this, believing that excessive bleeding is det- tendrils (Hegwood et al., 1983; Ferree et al., earliest and have the highest quality. The de- rimental. He collected as much as 1.5 kg of 1983; Poling et al., 1987; U.S. Dept. of Ag- gree of secondary fruit development is affected sap per vine over 2 months from vines pruned riculture, 1973). by cultivar, vine age, management practices, 20 Mar. Late pruning has been reported to New shoot growth and fruiting wood must and environmental conditions. delay shoot and berry development and reduce be prevented from developing on the per- Fruit development. Muscadine berries ap- yields (Newman, 1907; Onokpise and Inyang, manent trunk and arms leading up to the trel- pear to develop in the same manner as Euvitis 1987), but others found no decrease in yield lis wires, particularly for efficient mechanical grapes, following a double-sigmoidal growth with late pruning (Dearing, 1938; Loomis, harvest. An exception is the occasional need curve (Carroll, 1985; Pratt, 1971). Sugar con- 1943). Dearing (1938) concluded that pruning to train replacement trunks or cordons. Dear- tent and pH increase and acid content de- should be confined to the period between mid- ing (1938) recommended regular, systematic creases as berries mature (Carroll, 1985). winter and bud swell. Increased susceptibility replacement of cordons over an 8-year pe- Although fruit size at maturity differs consid- to winter injury has been noted following riod. New shoot growth is easily rubbed off erably with cultivar, perfect-flowered types tend pruning before mid-winter (Dearing, 1938). by hand in the spring and early summer. to be smaller (average 4.7 g/berry, ranging Early publications recommended pruning Naphthaleneacetic acid in white latex paint from 2.2 to 7.4 g/berry) than pistillate-flow- muscadines to short spur systems on per- has been used successfully to prevent shoot ered types (average 5.8 g/berry, ranging from manent cordons. (Armstrong et al., 1934; development on established trunks, while 2.8 to 15.0 g/berry) Andersen et al., 1985; Murphy et al., 1938; Savage et al., 1941), physical wraps prevented shoot development Balerdi and Mortensen, 1969; Carroll, 1985; but supporting data were lacking. Loomis but induced root development on the trunk Hegwood et al., 1983; Moore and Bowden, (1943) found higher yields in a 3-year trial (Takeda et al., 1983). 1976; Poling et al., 1985; Williams, 1954). when mature ‘Scuppernong’ and ‘Thomas’ Pollination and fruit set. Small vineyards of Muscadine berries form an abscission zone vines were pruned to four-bud spurs in com- female cultivars might depend on male vines between the fruit and rachis (Sherman, 1963; parison to pruning to a six-cane Kniffen sys- in adjacent woods, but commercial production Mitchell, 1979). In most cultivars, the berry tem with 2.3-m canes. In a 5-year study, of such cultivars requires planting male vines matures before this abscission layer is fully there was no effect on yield when mature every third vine in every third row (Husmann formed, although there is variation among cul-

734 HORTSCIENCE, VOL. 25(7), JULY 1990 eastern United States were reported to average ing and methods (Harkness et al., tivars and years (Sherman, 1963). Fruit re- -1 moval force has been positively correlated with 4.4 to 9.0 t·ha in 1973 (U.S. Dept. of 1987). Effects of temperature during press- Agriculture, 1973). North Carolina reported ing and pressing aids (e.g., rice hulls) are juice soluble solids content in ‘Carlos', ‘Mag- -1 nolia’, ‘Tarheel’, and ‘Noble’ (Mitchell, 1979). yields of 7.8 t·ha as recently as 1982 (Pol- being evaluated as a means to increase mus- ing, 1982). However, many growers are cadine juice yield while maintaining juice When harvested, immature fruit abscission -1 zones tend to tear, resulting in a “wet stem achieving regular yields of 22 t·ha . Yields quality (Harkness et al., 1987; Sims et al., scar”, tom skin, and leaking juice. Fruit in in published cultivar trials vary widely, but 1988). better-yielding cultivars generally range from Vineyard floor management. Herbicide weed this condition deteriorate and decay rapidly. -1 ‘Carlos’ is the best cultivar developed to date 11 to 34 t·ha at a vine spacing of 3.7 × control in the row with grass alleys between for stem scar rating, usually 10% wet scar at 6.0 m (e.g., Andersen et al., 1985; Balerdi rows has become standard practice, but bare harvest, and is often used as the standard for and Mortensen, 1969; Goldy, 1988; Heg- soil can create problems in highly erodible sites. evaluating new cultivars (Lane, 1977a, 1980; wood et al., 1983; Moore and Bowden, 1976). Early muscadine bulletins recommended in- Yields of ‘Doreen’ have been recorded as tercropping “hoed” crops such as melons, to- Lane and Bates, 1987; Mitchell, 1979). Other -1 cultivars with high dry stem scar ratings are high as 40 t·ha in cultivar trials (Hegwood matoes, peanuts, cotton, tobacco, and legume ‘Golden Isles’ (Lane and Bates, 1987), et al., 1983). crops to add income, especially during the 3 ‘Southland’ (Hegwood et al. 1983), ‘Summit’ Fruit quality and juice yield. Due to un- to 5 years before significant ‘yields were ob- (Lane, 1977a), ‘Triumph’ (Lane, 1980), and even ripening, methods of fruit sorting and tained from the muscadine vines (Husmann ‘Thomas’ (Sherman, 1963). ‘Tarheel’ tends to grading would be particularly advantageous and Dearing, 1916). Cereals and corn were not be intermediate in stem scar rating (Mitchell, for fresh market sales. Successful grading recommended. Frequent cultivation has been 1979). into ripeness categories has been accom- used to develop a deep root-system, but this Harvest. With good vineyard management, plished by a reflected light spectrophoto- practice is no longer recommended because muscadines will produce a commercial yield meter (Ballinger et al., 1978) and by density muscadines tend to be very shallow-rooted, in the third year after planting and mature yields separations in graded brine solutions (Lanier even in well-drained soils. Current research on by the 6th to 7th year (Dearing, 1938). Be- and Morris, 1979). intercropping is being conducted to aid small cause of the tendency to shatter (abscise), Factors affecting quality of juice, wine, farmers (O. Bandele, personal communica- muscadines are usually harvested as single and fresh use, and yields of juice per weight tion). Interplanted clover is also being ex- berries (Mitchell, 1979). However, whole of berries, were recently reviewed by Carroll plored as a mulch and N source (C.P. Hegwood, clusters of cultivars with less tendency to shat- (1985). Muscadine juice tends to be lower personal communication). ter, such as ‘Fry’ and ‘Pride’, may be clipped in soluble solids concentration (SSC) and ti- Irrigation and fertility. Irrigation has been by hand for the fresh market (Lane, 1972). tratable acidity than V. labruscana and V. recommended only recently in the mid-Gulf Traditionally, muscadines are harvested semi- vinifera juice (Carroll, 1985). Sugar and acid of Mexico region (Hegwood et al., 1983), mechanically. The trellis and vine arms are frequently must be added to the juice to pro- and is not common in the Carolinas and beaten with baseball bats, shaking the fruit off duce a wine with sufficient acid and alcohol Georgia (Ferree et al., 1983, Poling et al., the vine onto-a tarp or canvas frame (Clark, content. Carroll (1985) listed major juice 1987). Austin and Bondari (1988) found 1981; U.S. Dept. of Agriculture, 1973). This components and average concentrations for considerable variation in yield response of method can damage the vine and fruit. Fruit muscadine cultivars in current production as mature ‘Hunt’ vines to drip-irrigation over 8 bruised and covered with leaking juice due to fructose (5.51%), glucose (5.16%), sucrose treatment years, and a significant interaction harvest injury must be processed or sold fresh (1.89%), malic acid (0.50%), and tartaric with fertility level. Yields of nonirrigated within a few days. Most mechanical harvesters acid (0.36%). Soluble solids concentration vines fertilized annually with 1.4 kg/plant of operate in a similar manner to the semime- averaged 13.2% (10% to 18%), titratable 10N-4P-8K fertilizer were not significantly chanical method and cause similar problems. acidity (expressed as tartrate equivalent) different from vines irrigated to maintain soil A new mechanical harvester for muscadines is 0.84% (0.39% to 1.5%), and pH 3.14 (2.9 water potential > 15 kPa at 15- and 45-cm under development at Mississippi State Univ. to 3.4). As with other American grapes, the depths and fertilized with 0.9 kg/plant of the This harvester vibrates the vine arms, rather significant sucrose content of muscadine juice same fertilizer. Irrigation markedly reduced than beating them, and shows promise for both is in contrast to the trace concentrations of alternate bearing. processing and fresh-market muscadines. (2- sucrose in V. vinifera grapes. Grower con- Where used, irrigation is almost exclu- Chloroethyl)phosphonic acid (ethephon) has tracts for sale of muscadines for juice pro- sively applied through drip systems. Irriga- been used successfully as an aid to mechanical duction specify SSC of 13% to 15%, with tion is particularly important during the first and semimechanical harvest of muscadines by premiums paid for SSC up to 18% (R.T. 2, years to hasten vine establishment and promoting development of the fruit abscission Kahrer, personal communication). No men- training (Hegwood et al., 1983). Bearing layer (Lane and Flora, 1979; Mortensen, 1980; tion is made of juice pH or titratable acidity. vineyards should be well-irrigated during the Phatak et al., 1980). Flavor components are more highly corre- first part of the season to support shoot Average harvest date across all cultivars lated with juice pH than with titratable acid- growth, flowering, and initial fruit set. How- ranges from 24 Aug. in central Florida to 15 ity or SSC (R.P. Vine, personal com- ever, after set of the primary crop, irrigation Sept. in Mississippi, Georgia, and the Car- munication). Carroll (1985) stated that total should be limited during the remainder of the olinas (C.P. Hegwood, R.P. Lane, and J.A. acidity and pH are probably more critical season to reduce secondary fruit set and un- Mortensen, personal communications). The than sugar content as a harvest index for wine even ripening and to reduce late-season shoot period from flowering to harvest is fairly production. He recommends harvesting at a growth, delayed dormancy,and susceptibil- constant across production areas, varying from total acidity above 0.5% and pH of 3.0 to ity to winter injury. 106 to 113 days. Individual cultivars vary 3.4. Bates et al. (1986) recommend musca- Fertilizer recommendations for musca- considerably in average harvest date, pro- dine harvest for wine before full maturity dines have been largely based on bunch grape viding a harvest season of »30 days. with juice pH of 3.0 to 3.5. Harvesting by practices (Hegwood et al., 1983; Ferree et Fruit yield. -Early in this century, yields pH alone in the range of pH 3.1 to 3.3 has al., 1983; Poling et al., 1987; U.S. Dept. of of mature muscadine vines averaged 6.7 to been recommended for unfermented juice Agriculture, 1973). Muscadine fertility re- 10.1 t·ha-1, and ranged from 4.4 to 17.9 (R.P. Vine, personal communication). search prior to 1966 was largely concerned t·ha-1 depending on site, season, care, and Low juice yields per tonne of fruit due to with Mg deficiency (Cook, 1966). Much of cultivar (Husmann and Dearing, 1916). thick skins and mucilaginous pulp is a sec- the muscadine production is on sandy soils ‘Scuppernong’ was noted to produce low ond problem in processing muscadines with low cation exchange capacity and low yields, averaging 5 t·ha-1. Despite changes (Harkness et al., 1987; Sims et al., 1988). Mg levels. Muscadines are particularly prone in cultivars and management practices, the Juice yields of muscadines range from 46% to Mg deficiency, and symptoms of inter- average commercial yield has remained rel- (w/w) to 54%, as compared to 67% to 75% veinal leaf chlorosis are frequently apparent atively unchanged. Yields over the south- from ‘Concord’, using conventional crush- from the middle to end of the growing season

HORTSCIENCE, VOL. 25(7), JULY 1990 735 (Lott, 1948; Hagler, 1949). The problem can vine death in some cases (Hopkins et al., boom lasted until 1926, when grape prices be corrected by soil amendment, foliar ap- 1974). The causal organism of crown gall dropped due to over-production, which was plication, or trunk injection of MgSO4 (Lott, [Agrobacterium tumefaciens (E.F. Smith & followed by the Great Depression, when many 1948, 1952), but best results were obtained Townsend) Conn. Biovar. 3] has been found vineyards were abandoned. National prohi- by foliar sprays of MgCl2 and Mg(NO3)2 as a latent, systemic infestation in nearly all bition ended in 1932 with adoption of the (Hagler, 1957). Apparent yield responses to muscadines vines tested (Griffin and Graves, 21st Constitutional Amendment, but prohi- B application have been noted in Georgia, 1988). However, actual gall formation and bition forces remained strong in the South (Boswell et al., 1980). Ragland (1940) noted associated vine injury or death are generally and the muscadine industry did not rebuild. response to potash, but not nitrate or P, in only induced after winter injury and are not In 1935, the Federal Emergency Relief reversing symptoms of leaf yellowing and frequently observed. The disease is most apt Administration began a program to promote scorch in a mature muscadine vineyard. A to be a problem where low winter tempera- muscadine fruit and wine production as an series of papers by Cummings and co-work- tures are limiting. No practical controls exist aid to Southern families on Rural Relief and ers in North Carolina reported considerable for Pierce’s disease or crown gall. Virus dis- to agronomic growers facing economic dis- variation in leaf and fruit nutrient content eases have not been reported in muscadines, aster with cotton, tobacco, and other crops through the season and between years (Cum- but it is likely that they occur. (Gohdes, 1982). The plan called for planting mings and Lilly, 1984, and papers cited Insect control is primarily limited to grape 2 million vines. The problems of mass prop- therein). Optimum yields occurred over soil root borer (Vitacea polistiformis Harris) (Dutcher agation on this scale were described by pH values of 5.5 to 7.0, and yield was in- et al., 1988; McGiffen and Neunzig, 1985). Woodroof (1936). Some of these grapes were creased at all pH levels by increasing annual Control of this pest is extremely difficult be- planted and brought into production; how- N application from 50 to 83 kg·ha-1 (Cum- cause of the subterranean feeding of larvae on ever, the program as a whole-was not suc- mings and Lilly, 1984). roots and the lack of distinct early symptoms cessful and was strongly opposed by Diseases and pests. Muscadines have a (All et al., 1987). Early claims that musca- prohibition forces. high degree of resistance to pests and dis- dines were immune to grape root borer have Significant expansion of muscadine acreage eases, especially Pierce’s disease (Xylella been shown to be erroneous. No complete occurred in the early 1960s and continued fastidiosa Wells et al.), which severely lim- control strategy has been developed, but par- through the early 1980s with increased na- its production of Euvitis species in the south- tial control is achieved with a soil drench of tional consumption of wine (Brooks, 1978; eastern United States (Bouquet., 1980; chlorpyrifos insecticide (Lorsban-4E, Dow, Gohdes, 1982; Poling et al., 1987). Again, Hedrick, 1908). Vitis rotundifolia is among Midland, Mich.) around the trunk, in combi- muscadine production, principally for wine, the most resistant grape species to the grape nation with cultural practices such as weed was seen as a means to aid depressed state insect (Daktulosphaira vitifolia control and temporary soil mounding (All et economies and as a profitable alternative crop Fitch), which causes devastating root injury al., 1987). Pierce’s disease is vectored by for agronomic growers. In 1965, North Car- to V. vinifera (Husmann and Dearing, 1916). sharpshooter leafhoppers (family Cicadelli- olina appropriated funds to stimulate re- Muscadine pests noted early in this century dae, subfamily Tettigelinae), but control of these search on breeding, production, and marketing were black rot [Guignardia bidwellii (Ellis) insects is difficult (Pearson and Goheen, 1988). of muscadines (Gohdes, 1982). Muscadine Viala & Ravaz] and grapevine flea beetle Other pests, such as mites, aphids, grape flea vineyards in North Carolina increased from (Altica chalybea Illiger) (Husmann and beetle (Altica chlybea Ill.), grape curculio 240 ha in 1968 to >1000 ha in 1976. In Dearing, 1916). As muscadines became more (Craponius inaequalis Say), and grape berry 1978, muscadine production in Georgia and extensively planted, additional pests were moth (Endopiza viteana Clemens) are occa- included 300 and 200 ha, re- recognized. Excellent descriptions of mus- sional problems and are controlled only if sig- spectively, with less production in , cadine diseases and pests are contained in nificant injury is observed (Hegwood et al., , Florida, and Mississippi (Brooks, references by Pearson and Goheen (1988), 1983; Ferree et al., 1983; Poling et al., 1987; 1978). Major commercial use of muscadines McGiffen and Neunzig (1985), and Dutcher Savage, 1941). remained wine production, but fresh fruit sales et al. (1989). were increasing in importance. As recently Primary disease problems of muscadines HISTORY OF PRODUCTION as the early 1970s production in Georgia are fruit rots, especially black rot, bitter rot AND USE and the Carolinas depended on the old cul- [Greeneria uvicola (Berk. & Curt.) Puni- The muscadine industry has experienced tivars Hunt, Scuppernong, and Thomas (U.S. thalingam, syn. Melanconium fuligineum fluctuations closely tied to sales of musca- Dept. of Agriculture, 1973). With expansion (Scribner & Viala) Cav.], ripe rot [Colleto- dine wine (Gohdes, 1982; Morton, 1988; of the industry, the leading cultivars in North trichum gloeosporioides (Penz.) Penz. & Reimer, 1909). In 1810, Washington County, Carolina shifted by 1976 to ‘Carlos’ (47%), Sacc., teleomorph Glomerella cingulata N.C., produced 5176 liter (1368 gallons) of ‘Magnolia’ (20%), and ‘Scuppernong’ (12%), (Stonem.) Spauld. & Schrenk], and macro- wine from native grapes, most of it from followed by ‘Fry’, ‘Higgins’, ‘Hunt’, and phoma rot [Botryosphaeria dothidea (Moug. ‘Scuppernong’ (Reimer, 1909). The industry ‘Noble’ (Gohdes, 1982). Prices paid for fruit ex Fr.) Ces. & de Not., anamorph Macro- continued to grow, with vineyards in North by declined and muscadine produc- phoma sp.] (Ferree et al., 1983; Hegwood Carolina as large as 243 ha (Reimer, 1909). tion in North Carolina fell to 688 ha by 1981 et al., 1983; Jabco et al. 1985; Poling et al., Until national prohibition, »60% of total (Poling et al., 1987). The important cultivars 1987; Savage, 1941). If severe, angular leaf muscadine production was used for wine in North Carolina in 1985 were the same as spot (Mycosphaerella angulata Jenkins) may (Gohdes, 1982; Husmann and Dearing, 1916). in 1976, but ‘Magnolia’, and especially cause leaf abscission, resulting in smaller “ Dare”, produced by Paul Garrett, ‘Scuppernong’, were decreasing in impor- berries and lower sugar content than from was the most popular wine in the United States tance (Poling et al., 1985). Carroll (1985) healthy vines. Powdery mildew [Uncinula before prohibition (Gohdes, 1982). This wine recently estimated that bronze cultivars grown necator (Schw.) Burr.] has been reported as was initially pure muscadine, but later be- for wine made up 96% of North Carolina a problem on .‘Magnolia’, ‘Regale’, and came a blend of muscadine and California production. In 1980, fresh fruit was reported ‘Tarheel’ in Georgia and North Carolina V. vinifera. The influence of Garrett on the as the primary use for muscadines in Geor- (Lane, 19776; Poling et al., 1985). These muscadine industry is described by Gohdes gia, based largely on ‘Fry’ (Phatak et al., diseases have been controlled with cultural (1982). Adoption of prohibition, first at the 1980). Recent development of a major juice practices and fungicides, although the num- state level, and then nationally in 1919 with plant in Mississippi has provided an addi- ber of fungicides registered for use on mus- passage of the Volstead Act and the 18th tional stimulus to the muscadine industry. cadines has decreased in recent years. Constitutional Amendment, brought an end Production in Mississippi increased from 45 Muscadines are more resistant to Pierce’s to commercial wine production (Gohdes, ha in 1984, primarily for small wineries, to disease (Xyella fastidiosa Wells et al.) than 1982). However, increased demand for home »340 ha in 1988 (G. Feltenstein, personal Euvitis grapes, but the disease has been noted wine making actually led to a boom in both communication). Expansion of muscadine in muscadines and has been associated with East and West Coast grape production. The vineyards is continuing in Mississippi and is

736 HORTSCIENCE, VOL. 25(7), JULY 1990 expected to exceed 500 ha in 1990. Total SURVEY AND CURRENT STATUS OF based on the number of states listing each grape production (bunch plus muscadine) in THE MUSCADINE INDUSTRY cultivar as important (Table 2). Currently, Arkansas, Georgia, North Carolina, South I conducted a survey in 1988 to gain a the two most important cultivars are ‘Carlos’ Carolina, and Missouri averaged 14,800 t/ more detailed picture of the bunch grape and and ‘Magnolia’ (Group A). Nine additional year during 1984 through 1986 (U.S. Dept. muscadine grape industries in the southeast- cultivars also ranked highly, being listed in of Agriculture, 1987), which represented ern United States, Experiment station re- five to six states (Group B) or three to four 0.3% of total U.S. production, or 11% of searchers and extension specialists dealing states (Group C). The 14 cultivars in Group grapes produced in Eastern states. with grapes in 15 states (Table 1) were con- D were listed in commercial production in tacted and asked to provide the following one or two states. Not listed are many cul- information for each state: 1) An estimate of tivars that are in minor commercial produc- Table 1. Current muscadine production in 15 the number of hectares producing muscadine tion or are grown in backyards. Many newly southeastern states.z and bunch grapes; 2) the most common mus- released cultivars are being planted and will cadine cultivars grown; 3) primary uses and change cultivar ranking in the future. ‘Do- markets for grapes; and 4) primary problems reen’ is a new cultivar increasing in impor- of the muscadine and bunch grape industries. tance, while ‘Hunt’, ‘Magnolia’, and Muscadine hectarage in the southeastern ‘Scuppernong’ are examples of important United States is estimated at > 1600 ha, 25% older cultivars that are being replaced. ‘Fry’ of the total Vitis hectarage in these states. remains the primary cultivar for fresh-market Interior southeastern states have more than production. Current expansion of muscadine twice the hectarage of all grapes as that in production in Mississippi is based heavily on coastal states. Muscadine production is con- ‘Carlos’ and ‘Doreen’, but many other cul- centrated in the coastal states, accounting for tivars are also being planted. Primary char- 2% of the total grape hectarage in interior acteristics and uses of the more-important or states and 76% in coastal states. States with well-known cultivars are summarized in Ta- > 200 ha of muscadine grapes were (most to ble 3. Current problems and strengths of the least) Georgia, Mississippi, North Carolina, muscadine-industry are summarized in Table Florida, and South Carolina. Major bunch 4. Many of the environmental and marketing grape-producing states in the South are Texas, problems listed are also factors limiting bunch Arkansas, Missouri, and Virginia, with ma- grape production. jor emphasis on V. vinifera for wine pro- Commercial interests continue to demand duction. In general, muscadines are grown improved cultivars and cultural methods for where Pierce’s disease is limiting and Amer- muscadine production. Cultural methods de- ican and French-American hybrid Euvitis veloped for bunch grapes often cannot be di- grapes are grown where cold injury is lim- rectly applied to muscadines due to marked iting. Vitis vinifera is preferred because of genetic differences between Muscadinia and greater profitability in wine production and Euvitis. Continued commercial development is grown where neither Pierce’s disease nor depends heavily on research to improve pro- cold injury is limiting. duction and processing methods, and to de- A quantitative comparison of relative cul- velop superior cultivars for fresh market and tivar importance is not possible because for processing. Work with Muscadinia x Eu- zInformation based on responses of muscadine re- muscadine production and hectarage statis- vitis hybrids will also be important in devel- searchers and extension specialists surveyed in each tics for the region are not available. How- oping new rootstock and fruiting cultivars for state. ever, cultivars were grouped qualitatively, both bunch grape and muscadine production.

Table 2. Primary muscadine cultivars grown by state.z

zInformation based on responses of muscadine researchers and extension specialists surveyed in each state. Group indicates approx- imate overall cultivar importance. Cultivar importance in each state is estimated as primary (*) or secondary (+).

HORTSCIENCE, VOL. 25(7), JULY 1990 737 Table 3. Characteristics and primary uses of important or well-known muscadine cultivars.

zFlower type is perfect (Per) or female (Fern). yFruit color is bronze (Brz), purple (Pur), or black (Blk). xFruit size is based on average berry fresh weight: Small (2.1 to 4.0 g), Med (4.1 to 6.0 g), Lrg (6.1 to 8.0 g), VLrg (8.1 to 10.0 g). wHarvest period is Early (25 Aug.-9 Sept.), Mid-season (10-24 Sept.), or Late (25 Sept.-5 Oct.). vUse is categorized as primary (*) or secondary (+).

Literature Cited Table 4. Problems and strengths of the muscadine industry. All, J.N., J.D. Dutcher, and M.C. Saunders. 1987. Con- Problem Strength trol program for the grape root borer in grape vineyards of the eastern United States. Down to Earth 43:10-12. Environmental Andersen, P.C. and B.V. Brodbeck. 1988. Water and Cold injury in northern range of production. Relatively drought-tolerant. solute fluxes through pruned muscadine grape spurs. Poor tolerance to wet or heavy soil. HortScience 23:978-980. Mg deficiency common. Andersen, P.C., M.W. Byran, and L.H. Baker. 1985. Pests, diseases Effect of two wire vertical and Geneva Double Curtain Fruit rots. training systems on berry quality and yield of mus- Broad tolerance to diseases and pests compared cadine grapes. Proc. Fla. State Hort. Soc. 98:175- Grape root borer. to Euvitis grapes. 178. Physiological, cultural, genetic Andrew, C.P. 1981. Effect of trellis on yield of mus- Uneven fruit ripening. Will survive neglect, long life in good sites. cadine grape cultivars. HortScience 16:422. (Abstr.) “Wet stem scar”, rapid fruit decay. Potential for very high yields with good man- Armstrong, W.D. 1936. New varieties and pollination of muscadine grapes. Proc. Amer. Soc. Hort.. Sci. 33:450- Poor fruit set due to “cap sticking” as flowers agement and cultivar choice. 452. open, especially ‘Sugargate’ and ‘Fry’. Improved cultivars are being developed, - Armstrong, W.D., T.A. Pickett, and M.M. Murphy, Jr. Excessive shoot growth/fruit load. less cultivars may be possible. 1934. Muscadine grapes-culture, varieties, and some Poor plant establishment and early growth. Improved methods for wine and processed properties of juice. Ga. Expt. Sta. Bul. 185. Low yields due to poor cultivars, site, or vine- products developed for flavor and color sta- Austin, ME. and K. Bondari. 1988. A study of cultural yard management. bility. and environmental factors on the yield of Vitis rotun- difolia. Scientia Hort.. 34:219-227. Economic Austin, M. E. and K. Bondari. 1989. Yield of muscadine Unstable markets. Good current market diversity in fresh mar- grape cultivar Hunt as influenced by plant and row Low prices paid by processors. ket, wine, juice, and processing industries. spacings and position of the male plant. Applied Agr. Poor consumer familiarity with muscadines out- Incentive programs have been established by Res. 4:1-9. side of the southeastern United States. states to help finance costs of vineyard estab- Bailey, L.H. 1937. The standard encyclopedia of horti- Large initial investment. lishment. culture. Macmillan, New York. Significant yield not obtained until 3 to 5 years Balerdi, C.F. and J.A. Mortensen. 1969. Performance of muscadine grapes (Vitis rotundifolia Mich.) in central after planting. Florida. HortScience 4:252-253. Ballinger, W.E., E.P. Maness, A.J. Makus, and D.E. Hort. Sci. 100:58-60. Davidis, U.X. and H.P. Olmo. 1964. The Vitis vinifera Carroll, Jr. 1974. A comparison of and Brightwell, W.T. and M.E. Austin. 1975b. Influence of x V. rotundifolia hybrids as phylloxera resistant root- wine color quality in black grapes of 39 clones of Vitis plant spacing on yield of muscadine grape. J. Amer. stocks. Vitis 4:129-143. rotundifolia Michx. J. Amer. Soc. Hort. Sci. 99:338- Soc. Hort. Sci. 100:374-376. Dearing, C. 1917. Muscadine grape breeding. J. Hered. 341. Brooks, J.F. 1978. Muscadine grapes in the Southeast. 8:409-424. Ballinger, W.E., W.F. McClure, W.B. Ncsbitt, and E.P. HortScience 13:264. (Abstr.) Dearing, C. 1938. Muscadine grapes. U.S. Dept. of Agr. Maness. 1978. Light sorting muscadine grapes (Vitis Carroll, D.E. 1985. Muscadine manes: Factors influenc- Farmers’ Bul. 1785. rotundifolia Michx.) for ripeness. J. Amer. Soc. Hort. ing product quality, p. 177-197.In: H.E. Pattee (ed.). Dearing, C. 1947. Muscadine grapes. U.S. Dept. of Agr. Sci. 103:629-634. Evaluation of quality of fruits and vegetables. AVI, Farmers’ Bul. 1785. Bates, R.P., J.A. Mortensen, and T.E. Cracker. 1980. Westport, Conn. Dearing, C. 1948. New muscadine grapes. U.S. Dept. Florida grapes: The next decade. Proc. Fla. State Hort. Chaparro, J.X., R.G. Goldy, B.D. Mowrey, and D.J. of Agr. Circ. 769. Soc. 93:120-124: Werner. 1989. Identification of Vitis vinifera L. x Detjen, L.R. 1917a. Inheritance of sex in Vitis rotundi- Bates, R.P., T. Tejada, and J.A. Mortensen. 1986. Ef- Muscadinia latundifolia Small hybrids by starch gel folia. North Carolina Agr. Expt. Sta. Tech. Bul. 12. fect of bunch and muscadine grape maturity on fin- electrophoresis. HortScience 24:128-130. Detjen, L.R. 1917b. Pollination of the rotundifolia grapes. ished wine. Proc. Fla. State Hort.. Soc. 99:194-200. Clark, J.R. 1981. Catch frames for harvesting musca- J. El. Mitchell Sci. Soc. 33:121-127. Boswell, F.C., R.P. Lane, and K. Ohki. 1980. Field dines. Mississippi Agr. & For. Expt. Sta. Info. Sheet Dutcher, J.D., K.C. McGiffen, and J.N. All. 1988. En- studies with boron on muscadine grapes. Commun. 1306. tomology and horticulture of muscadine grapes, p. 73- Soil Sci. Plant Anal. 11:201-207. Cook, J.A. 1966. Grape nutrition, p. 777-812. In: N.F. 90. In: M.K. Harris and C.E. Rogers (eds.). The en- Bouquet, A. 1980. Vitis x Muscadinia hybridization: A Childers (ed.). Temperate to tropical fruit nutrition. tomology of indigenous and naturalized systems in ag new way in grape breeding for disease resistance in Horticultural Publ., Rutgers Univ., New Brunswick, riculture. Westview Press, Boulder, Colo. France. Proc. 3rd Intl. Symp. on Grape Breeding. p. N.J. Einset, J. and C. Pratt. 1975. Grapes, p. 130-153. In: 42-61. Cummings, G.A. and P. Lilly. 1984. Soil pH rate for J. Janick and J.N. Moore (eds.). Advances in fruit Brightwell, W.T. and M.E. Austin. 1975a. Influence of fruit yield and elemental concentration of muscadine breeding. Purdue Univ. Press, West Lafayette, Ind. trellis type on yield of muscadine grape. J. Amer. Soc. grapes. HortScience 19:831-832. Fennell, J.L. 1940. Two North American species of Vitis.

738 HORTSCIENCE, VOL. 25(7), JULY 1990 J. Wash. Acad. Sci. 30:15-19. muscadine grape for wine. Georgia Agr. Expt. Sta., of muscadine and bunch grape cultivars in North Flor- Ferree, M.E., R.M. Crassweller, and G.W. Krewer. 1983. Res. Rpt. 520. ida. Proc. Fla. State Hort. Soc. 100:310-312. Commercial muscadine grape culture. Georgia Coop. Lane, R.P. and L.F. Flora. 1979. Effect of ethephon on Pearson, R.C. and A.C. Goheen. 1988. Compendium of Ext. Serv. Bul. 739. ripening of ‘Cowart’ muscadine grapes. HortScience grape diseases. APS Press, St. Paul, Minn. Gohdes, C. 1982. Scuppernong, North Carolina’s grape 14:727-729. Phatak, S.C., M.E. Austin, and J.S. Mason. 1980. Ethe- and its . Duke Univ., Durham, N.C. Lanier, M.R. and J.R. Morris. 1979. Evaluation of den- phon as harvest-aid for muscadine grapes. HortScience Goldy, R.G. 1988. Variation in some yield determining sity separation for defining fruit maturities and matu- 15:267-268. components in muscadine grapes and their correlation ration rates of once-over harvested muscadine grapes. Poling, B. 1982. A troubled muscadine industry. Facts to yield. Euphytica 39:39-42. J. Amer. Soc. Hort. Sci. 104:249-252. for farm markets. Summer. p. 4-6. N.C. Agr. Ext. Goldy, R., R. Emershad, D. Ramming, and J. Chaparro. Lee, N. and H.Y. Wetzstein. 1988. Protoplast isolation Ser. 1988. Embryo culture as a means of introgressing and callus production from leaves of tissue-cultured Poling, B., C.M. Mainland, and J.B. Earp. 1987. Mus- seedlessncss from Vitis vinifera to V. rotundifolia. Vitis-spp. Plant Cell Rpt. 7:531-534. cadine grape production guide for North Carolina. N.C. HortScience 23:886-889. Lee, N. and H.Y. Wetzstein. 1990. In vitro propagation Agr. Ext. Ser. Bul. AG-94. of muscadine grape by axillary shoot proliferation. J. Goldy, R.G., E.P. Maness, H.D. Stiles, J.R. Clark, and Pratt, C. 1971. Reproductive anatomy in cultivated Amer. Soc. Hort. Sci. 115:324-329. M.A. Wilson. 1989. Pigment quantity and quality grapes-A review. Amer. J. Enol. Viticult. 22:92- characteristics of some native Vitis rotundifolia Michx. Liberty Hyde Bailey Hortorium. 1976. Hortus third: A 109. concise dictionary of cultivated in the United Amer. J. Enol. Viticult. 40:253-258. Ragland, C.H. 1940. Response of muscadine grapes to States and Cananda. 3rd ed. Macmillan, New York. Goode, D.K., Jr., G.W. Krewer, R.P. Lanf, and J.W. potassium and phosphorus fertilizers. Proc. Assn. Lider, L.A., H.P. Olmo, and A.C. Goheen. 1988a. Hy- Daniell. 1982. Rooting studies of dormant muscadine Southern Agr. Workers 41:154-155. brid grapevine rootstock. U.S. Pat. No. Plant 6,166. grape cuttings. HortScience 17:644-645. Reimer, F.C. 1909. Scuppernong and other muscadine Lider, L.A., H.P. Olmo, A.C. Gohcen. 1988b. Hybrid Goode, D.Z., Jr., and R.P. Lane. 1983. Rooting leafy grapes: Origin and importance. North Carolina Agr. grapevine rootstock named ‘VR 043-43’. U.S. Pat. muscadine grape cuttings. HortScience 18:944-946. Expt. Sta. Bul. 201. No. Plant 6,166. Gray, D.J. and L.C. Fisher. 1985. In vitro shoot prop- Schwartz, K. 1976. The origin and development of mus- Loomis, N.H. 1943. The influence of time and method agation of grape species, hybrids and cultivars. Proc. cadine grape varieties. Fruit Var. J. 30:90-92. of pruning on yields of muscadine grapes. Proc. Amer. Fla. State Hort. Soc. 98:172-174. Sherman, W.B. 1963. A morphological study of fruit Griffin, D.E. and C.H. Graves. 1988. Seasonal distri- Soc. Hort. Sci. 42:418-420. Loomis, N.H., M.M. Murphy, and F.F. Cowart. 1949. abscission of the muscadine grape, Vitis rotundifolia. bution of Agrobacterium spp. in Vitis rotundifolia vines. MS Thesis, Mississippi State Univ., Mississippi State. Phytopathology 78:627. (Abstr.) The effect of different methods of spur pruning upon the production and growth of muscadine grapes. Proc. Sims, C.A., R.P. Johnson, and R.P. Bates. 1988. Re- Griffin, D.E. and C.H. Graves. 1989. Micropropagation sponse of a hard-to-press Vitis rotundifolia cultivar and of muscadine and its use in the development of path- Amer. Soc. Hort. Sci. 54:183-185. Lott, W.L. 1948. Mg injection in muscadine grape vines. a hard-to-clarify Euvitis hybrid to commercial enzyme ogen-free propagation stock. Proc. Mississippi Assn. preparations. Amer. J. Enol. Viticult. 39:341-343. Plant Pathol. Nematol. p. 5-6. (Abstr.) Proc. Amer. Soc. Hort. Sci. 52:283-288. Lott, W.L. 1952. Mg deficiency in muscadine grape vines. Small, J.K. 1913. Flora of the southeastern United States. Hagler, T.B. 1957. Effect of magnesium sprays on mus- 2nd ed. (published by the author). cadinc grapes. Proc. Amer. Soc. Hort. Sci. 70:178- Proc. Amer. Soc. Hort. Sci. 60:123-131. Stucky, H.P. 1919. Work with Vitis rotundifolia, a spe- 182. Mainland, C.M., W.B. Nesbitt, and R.P. Rohrbach. 1982. cies of muscadine grapes. Georgia Expt. Sta. Bul. 133. Hagler, T.B. 1949. Nutrient-element deficiency symp- Effects of dormant season hedging of three muscadine toms of muscadine grapes in sand culture. Proc. Amer. grape cultivars on production, fruit quality and vine Sudarsono, J. and R.G. Goldy. 1988. Effect of some Soc. Hort. Sci. 53:247-252. condition. HortScience 17:501. (Abstr.) growth regulators on the in vitro culture of three Vitis Harkness, E.M., R.P. Vine, M.J. Coign, and J.R. Heitz. McGiffen, K.C. and H.H. Neunzig. 1985. A guide to rotundifolia cultivars. HortScience 23:757. (Abstr.) 1987. The effect of different must treatments on mus- the identification and biology of insects feeding on Syamal, N.B. and G.I. Patel. 1953. A wild species of cadine juice quantity-and quality. Amer. J. Enol. Vi- muscadine and bunch grapes in North Carolina. N.C. grape in India. Proc. Amer. Soc. Hort. Sci. 62:228- ticult., Meeting Rpt. p. 10. (Abstr.) Agr. Res. Ser. Bul. 470. 230. Hedrick, U.P. 1908. The grapes of New York. State of Mitchell, E.N. 1979. Correlation of the force required to Takeda, F., V. Drane, and M.S. Saunders. 1983. Sprout New York Dept. Agr. 15th Annu. Rpt. vol. 3, Part 2. pick Rotundifolia berries and their soluble solids con- control in muscadine grapes. HortScience 18:237-238. Hegwood, C.P., R.H. Mullenax, R.A. Haygood, T.S. tent. Arner. J. Enol. & Viticult. 30:135-138. U.S. Department of Agriculture. 1973. Muscadine grapes; Brook, and J.L. Peeples. 1983. Establishment and Moore, J.N. and H.L. Bowden. 1976. Muscadine grapes a fruit for the South. USDA Farmers’ Bul. 2157. maintenance of muscadine vineyards. Mississippi Coop. in east-central Arkansas. Arkansas Farm Rcs. 25:4. U.S. Department of Agriculture. 1987. Agricultural sta- Ext. Ser. Bul. 913. Mortensen, J.A. 1971. Breeding grapes for Central Flor- tistics 1987. U.S. Govt. Printing Office, Washington Hopkins, D.L., H.H. Mollenhauer, and J.A. Mortensen. ida. HortScience 6:7-11. D.C. 1974. Tolerance to Pierce’s Disease and the associated Mortenscn, J.A. 1980. Effects of ethephon on ease of Walker, M.A., J.A. Wolpert, E.P. Vilas, A.C. Goheen, Rickcttsia-like bacterium in muscadine grape. J. Amer. harvest of muscadine grapes. Proc. Fla. State Hort. and L.A. Lider. 1989. Resistant rootstock may control Soc. Hort. Sci. 99:436-439. Soc. 93:143-145. fanleaf degeneration of grapevines. Calif. Agr. 43:13- Husmann, G. and C. Dearing. 1916. Muscadine grapes. Morton, J.K. 1988. North Carolina’s viticulture history, 14. U.S. Dept. of Agr. Farmers’ Bul. 709. home of America’s 1st cultivated vines. Vinifera Wine Weaver, R.J. 1976. Grape growing. Wiley-Interscience, Jabco, J.P., W.B. Nesbitt, and D.J. Werner. 1985. Re- Growers J. 15:248-252. New York. sistance of various classes of grapes to the bunch and Munson, T.V. 1909. Foundations of American grape cul- Williams, C.F. 1923. Hybridization of Vitis rotundifolia. muscadine grape forms of black rot. J. Amer. Soc. ture. T.V. Munson & Son, Denison, Texas. Inheritance of anatomical stem characters. North Car- Hort. Sci. 110:762-765. Murphy, M.M, Jr., T.A. Pickett, and F.F. Cowart. 1938. olina Agr. Sta. Tech. Bul. 23. Lane, R.P. 1972. Recent muscadine grape releases from Muscadine grapes: Culture, varieties and some prop- Williams, C.F. 1954. Breeding perfect-flowered musca- Georgia. Fruit Var. J. 26:38-39. erties of juices. Georgia Expt. Sta. Bul. 199. dine grapes. Proc. Amer. Soc. Hort. Sci. 64:274-278. Lane, R.P. 1977a. ‘Summit’ muscadine grape. Hort- Newman, C.C. 1907. Rotundifolia grapes. South Caro- Winkler, A.J., J.A. Cook, W.M. Kliewer, and L.A. Lider. Science 12:588. lina Agr. Expt. Sta. Bul. 132. 1974. General viticulture. Univ. of Calif. Press, Lane, R.P. 1977b. Yield of young muscadine grapes as Olien, W.C. 1989. Methods of planting and vine man- Berkeley. affected by cane pruning. J. Amer. Soc. Hort. Sci. agement for rapid establishment of muscadine grapes Woodroof, J.G. 1934. Five strains of the Scuppernong 102:379-380. in sandy soil. Proc. Fourth Viniculture Short Course. variety of muscadine grapes. Proc. Amer. Soc. Hort. Lane, R.P. 1978. Bunch grape research in Georgia. Vin- p. 25-29. Mississippi State Univ. Mississippi State. Sci. 32:384-385. ifera Wine Growers J. 5:63-65. Olmo, H.P. 1986. The potential role of (vinifera x ro- Woodroof, J.G. 1936. Developments in growing mus- Lane, R.P. 1989. ‘Triumph’ muscadine grape. Hort- tundifolia) hybrids in grape variety improvement. Ex- cadine grapes in the South. Proc. Amer. Soc. Hort. Science 15:322. perientia 42:921-926. Sci. 33:447-449. Lane, R.P. and R.P. Bates. 1987. ‘Golden Isles’, a new Onokpise, O.U. and I.D. Inyang. 1987. Early pruning Young, W.J. 1920. Muscadine grapes: Culture and va- rieties. South Carolina Agr. Expt. Sta. Bul. 205.

HORTSCIENCE, VOL. 25(7), JULY 1990 739