SYSTEMATIC BIC MAIM' The Classification of Organisms with special reference to the grapevine

Dr. Stephen J. Krebs and Winery Technology Napa Valley College

"Systematic Biology" is a method of categorizing and naming organisms that was developed by the Swedish botanist Carolus Linnaeus (1707-1779). It consists of a hierarchical framework that makes it simple to understand the similarities and differences among all living creatures.

There are seven basic categories in the "Systematic Biology" framework, shown below in uppercase font:

KINGDOM PHYLUM Sub-Phylum CLASS Sub-Class ORDER Sub-Order Super Family FAMILY Sub-Family GENUS Variety Clone

Every organism is assigned to a group at each of these levels.

There are a total of six Kingdoms: , Animal, Fungal, Archaebacteria, Eubacteria, and Protista (protozoa and slime molds that are part-fungus, part-animal in nature). These are the most inclusive categories and large differences may be found among the members of each Kingdom. Thus, we do not learn very much about individual organisms at this level of classification.

Next, each Kingdom is sub-divided into several Phyla (Phylum, singular; Phyla, plural), also sometimes called Divisions. These are still very broad and inclusive categories, but for organisms to be grouped together in a Phylum, they must have more shared characteristics than at the Kingdom level.

Each Phylum is sub-divided into Classes. Then the Classes are further sub-divided into Orders. It is easy to see that the total number of categories is increasing rapidly as we work through the hierarchy. The amount of similarity among organisms placed in each category is also increasing.

In horticulture, it is the next level of detail, the Family, that really becomes helpful in understanding the relationships among . At this level, organisms placed together in the same Family must have many shared characteristics. For example, , the Grapevine Family, contains only flower- producing plants that grow as .

Families are sub-divided into Genera (Genus, singular; Genera, plural), and each Genus is sub-divided into Species. The combination of the Genus and Species names is called the binomial, which means "two names". The binomial is often referred to as the scientific name of an organism. The full binomial is needed to precisely identify a particular organism.

All grapevines are members of the Genus . Among the shared characteristics of all Vitis plants are the production of seeds in berries, similar leaf and shoot forms and the same general growth habit.

The high degree of similarity of all members of the Genus Vitis is reflected in the fact that Vitis plants are compatible reproductively, which means that they can inter-breed to produce hybrid off-spring. The new hybrid plant is made up of a mixture of traits from both the male and the female parents.

All Vitis plants are also graft-compatible, which means that viable physical unions of two separate Vitis species plant parts can be produced. This makes it possible to join a desirable fruit-bearing type and a useful type to produce a grafted . The genetic identities of the two plant parts do not intermingle, so each portion of the grafted vine retains its own unique traits.

The Genus Vitis contains approximately 60 different Species, all of which are readily recognized as grapevines. In many cases, the differences between two Species are very subtle. About 30 of these Vitis Species are native to America and about 30 are native to Asia.

The Species category is sub-divided into Variety to distinguish among variants within a single Species. In horticulture, these varieties are grown for agricultural or ornamental purposes and are referred to as cultivated varieties, also called . In the same way, varieties that occur in wild populations are referred to as biological varieties or biovars. Variations within a single Variety of grapevine are referred to as Clones. The differences among Clones of a variety may be quite small. The plants may look nearly identical in the field, with differences only becoming apparent after the fruit has been made into wine.

Of the Vitis Species native to Asia, by far the most important is , which means "wine-bearing grapevine". Its native range is in the Caucasus Mountains between the Black Sea and the Caspian Sea. It was domesticated very early in human history and was eventually spread throughout the temperate zones of both hemispheres. Almost all the familiar grape varieties grown worldwide are members of Vitis vinifera, including wine, raisin and table grape types.

Several of the American Vitis Species are used as , because only these Species are resistant to the grapevine root pest known as . The roots of the American Vitis Species are not severely damaged because they have co-evolved with this native North American insect over a very long period of time. All Asian Species, including Vitis vinifera, are quickly killed when phylloxera feeds on the root system. By grafting the Asian Vitis vinifera fruiting varieties onto phylloxera-resistant American Vitis rootstocks, grapevines can be grown successfully on soils that are infested with the insect.

A set of conventions has been established for written references to binomials, varieties and clones:

1. The first letter of a Genus name is always in uppercase font.

2. The Species name is always completely in lowercase font.

3. Both the Genus and Species name are used at each mention.

4. The binomial must be either written in italics (computer) or underlined (typewriter or hand writing).

5. The first mention of the binomial must be written out fully; in subsequent mentions, the first letter of the Genus and the full Species name are used:

Vitis riparia and V. rupeatris grow in America.

There are two exceptions. Always fully write out the Genus name when:

a) It is the first word of a sentence. b) There is more than one Genus cited which starts with the same letter. 6. The name is not written in italics, but each word in the name should start with an uppercase font letter:

Chardonnay, Cabernet Sauvignon, Zinfandel and so on.

In formal writing, the Genus and Species is listed, then the abbreviation "cv.", followed by the variety name:

Vitis vinifera cv. Merlot

When a cultivar name has a modifying word that indicates color or contains a preposition, those words are written entirely in lowercase font:

Vitis vinifera cv. Pinot noir Vitis vinifera cv. Muscadelle du Bordelais

In the popular press, wine grape variety names are often written entirely in lowercase, but this represents incorrect spelling of proper nouns.

7. When a specific clone of a cultivar is mentioned, its name appears in quotation marks following the cultivar name:

Vitis vinifera cv. Chardonnay "Wente"

If the clone is only designated by a number, it is written using the number symbol without quotation marks:

Vitis vinifera cv. Cabernet Sauvignon #8

8. In very formal usage, the last name of the person who first classified an organism will appear after the binomial, written in normal font, with or without parentheses:

Vitis rupestris (Scheele) or Scheele

For organisms that were originally classified by Linnaeus, only the initial L. is used, with or without parentheses:

Vitis vinifera (L.) or Vitis vinifera L.

If an organism has been reclassified, the new binomial and the person who reclassified it are listed first, followed by the original binomial and the name of the person who first classified the organism in parentheses, using the preceding format. 3 Early history this came as part of village settlement. of Vitis, Muscadinia has only three known species, The estimated 10,000 cultivars of the Old World are Cultivation of the wine grape was under way in the restricted to the southeastern USA and northeastern thought to derive from the single wild species, V. Near East as early as the fourth millennium B.c. Vitis, Muscadinia (Vitaceac) Mexico. The colonists of the Carolinas cultivated vinifera, of Middle Asia, still found from northeastern There is no evidence of any cultivation west of Greece M. rotundifolia directly and its domestication dates Afghanistan to the southern borders of the Black and until the first millennium B.C. The products of the from the latter part of the seventeenth century. Since Caspian Seas. Legend and tradition favour ancient vine were exported westwards from very early times, H. P. 01rno this species does not hybridize naturally with sym- Armenia as the home of the first grape and wine to be followed later by the practices peculiar to viti- patric species of Vitis, it represents an example of the University of Davis USA culture. Small refuge areas isolated by the glacial culture and by domesticated varieties (Hclback, 1959). domestication of a single species in situ. Though isola- epochs are found scattered in southern Europe, but The westward movement fanned out from Asia Minor ted in nature from Kris, M. rotundifolia can be hybrid- their role in domestication is questionable. Negrul and Greece, following the Phoenician sea routes. ized experimentally with V. tnnifera and the cross (1938) has proposed three principal groups of cultivars : During the Roman period, the spread of the vine was has been explored as a means of improving the occidentalis, the small-berried wine grapes of western associated with that of the Christian faith; wine is a disease resistance of vinifera and the fruit quality of Europe; oricntalis, the large oval-berried table grapes; necessary ingredient in the consecration of the Mass. rotundifolia. and pontica, intermediate types of Asia Minor and In the Middle Ages, the Catholic monasteries through- The cytogenetics of the F, (vinifiraxrotundifolia) eastern Europe. out Europe were the guardians of select vineyards. I Introduction with 39 somatic chromosomes and its backcross The fruit of wild vinifera (sylvestris) is palatable and A vineyard covered by the eruption of Vesuvius in The grapevine is a perennial, woody vine climbing by derivatives have been studied in some detail (Patel and the wine is of a quality comparable to that made from A.D. 79 illustrates the practices recommended in the coiled tendrils. As a cultivated plant it needs support Olmo, 1955). Unlike hybrids between Vitis species present cultivars. It was used in situ long before any early Roman agricultural manuals of that era (Jashem- and must be pruned to confine it to a manageable form (which arc fertile), the intergencric hybrids are highly settlement occurred. Domestication started when ski, 1973). The vine followed the main river valleys, and CO regulate fruitfulness. The fruit (a berry) is or completely sterile, though occasional viable seeds migratory nomads marked forest trees (usually poplar, the Danube, Rhone, Rhine, Tiber and Douro and, by juicy and rich in sugar (15-25%), in roughly equal are obtained in some combinations. At meiosis about pear, willow, plum or fig) that supported particularly A.D. 55, the northernmost vineyards were being proportions of dextrose and levulose. It is the com- 13 bivalents are formed, with univalcnts. The genomic fruitful vines. This was most often near watering established along the Moselle Valley in Germany. mercial source of tartaric acid and is rich in malic acid. formula of the F / hybrid is thus 13 R`Rv+7A+6B, in holes serving their herds. Sparing these vines was The vinifcra grape was introduced to the New World Cultivation of the crop is largely concentrated in which 13 chromosomes of vinifera and rotundifolia are associated with a spiritual taboo respected by other at the time of discovery and later accompanied prac- regions with a Mediterranean-type climate, with hot homologous enough to pair. The ancient basic chromo- tribes as well. As sedentary agriculture developed and tically all the Spanish and Portuguese voyages of dry summers and a cool rainy winter period. some numbers in the family are probably 5, 6 and 7. the mixed deciduous forest was cleared, fruit trees discovery and conquest (Fig. 86.1). The first recorded The world's vineyards occupy about 11 Mha. The 1'i:is species arc thus ancient secondary polyploids in- and vines were spared along boundary lines where introduction to the east coast of the principal product is wine, the mean annual output of volving three- basic scts in the combination (6+7)+6 irrigation ditches were developed and the vines were WAS in 1621 by the London Company but this was which, 1964-68, was 280.9 MM. About 10 per cent = 19. Muscadinia species, on the other hand, are (6+7) out of reach of grazing animals. Vineyards as such probably preceded by the Spanish landings in Florida. of this production enters international trade. The +7 = 20. Both have undergone diploidization to give developed later when they could be protected by high The most recent incursions of vinifera are in tropical biggest producers circle the Mediterranean, with Italy, regular bivalent pairing. mud walls from the ever-present sheep and goats, but countries; thus, in 1958, vinifera was introduced into France and Spain in the lead. The production of table The delimitation of species in Vitis has been grapes consumed as fresh fruit is about 6 Mt and the extremely difficult if not altogether artificiaL Taxo- Fig. 86.1 Evolutionary geography of the grape, Vitis vinifera. leading growers are Italy, Turkey, Bulgaria, the USA, nomists have been reduced to using such ephemeral Greece and Portugal. Production of raisins, largely characters as degree of hairiness of the young shoots sun-dried fruit of seedless cultivars, reached 0.9 Mt or leaves. Many species are sympatric with one or in 1970, the leading producers being the USA, Turkey, more others and are extremely variable. This vari- Greece and Australia. ability reaches its greatest expression in passing from uniform tropical to subtropical environments where 2 Cytotaxonomic background great differences in rainfall exist within short distances La vie (1970) has summarized the cytotaxonomy of the and isolated communities of vines become differentia- family. Vial contains about 60 species, but botanical ted. The populations represent ecospecies rather than knowledge is incomplete. This genus is unique species. All known Visit species can be easily crossed amongst the 12 recognized in the family Vitaceac experimentally and the F 1 hybrids are vigorous and in having 38 very small somatic chromosomes that fertile. Studies of natural populations indicate that regularly form 19 bivalents at meiosis. Wild grapes hybridization has occurred and continues. Even though can be divided into three geographical groups: species cannot be delimited on degree of genetic American, Middle Asian and Oriental. North America, isolation, the species classification remains useful especially the southeastern and Gulf region of the because of its practical value in separating norms of USA, is particularly rich in Vitis. Bailey (1934) lists variation that arc important in breeding. For example, 28 species, but does not include Mexico. riparia is highly resistant to phylloxera but vinifcra Most other related genera, including Muscadinia, quickly succumbs; and riparia roots readily from dor- have 2n = 2x = 40. Formerly classified as a section mant cuttings but aestivalis does not. Evolution of Crop Plants Edited by Longthan N. W. Simmonds ScD AICTA FRSE FIBiol Scientific & Edinburgh School of Agriculture Technical Edinburgh • Il tland such as Isabella, Herbemont, Concord and others which had been introduced as exotics years before, showed the Phillipines from California. mutation but are of different types and involve different some tolerance. The French government then sent other cultural methods will place new demands on As the vintfira grape was introduced into zones genetic backgrounds (Olmo and Baris, 1973). specialists to the United States to discover the best beyond its natural range it often hybridized with sources of resistance. From Missouri and other the breeder. The long generation time from seed to fruit (three to five years) could, perhaps, be shortened native Vies to produce new races better adapted to 4 Recent history areas of the midwest, thousands of cuttings and seeds local environments. Thus, new American hybrids Vitis species are dioecious or, occasionally, subdioe- were sent back to France for local selection. A few by the use of biochemical methods of selection at the earliest possible stage of seedling development. arose along the Atlantic seaboard as spontaneous cious when some male flowers transform to hermaph- proved tolerant, were widely propagated and utilized seedlings which were prized as new varieties. The rodites. The sexual type is determined by three alleles. as resistant rootstocks. Selections of riparia, rupestris 6 References Alexander, Concord and Delaware are examples. In This is a primitive type of sex determination in which and berlandieri proved most useful. From about 1880 Bailey, L. H. (1934). Vites peculiares ad Americana bore- the Caribbean islands and Venezuela, introduced gross differentiation of sex chromosomes has not onwards, interspecific hybrids were deliberately bred alem. Gent. Herb., 3, 149-244. innifera has introgressed with the tropical caribea to occurred (Negi and Olmo, 1971). The primitive as rootstocks. Thus berlandieri was the best adapted to Helbaek, H. (1959). Domestication of food plants in the produce new vigorous races of Criollas that are disease hermaphrodite is Su*Su*. A dominant mutation, calcareous soils but rooted with difficulty, so it was old world. Science, N.Y., 130, 365. resistant and tolerant of the climate, giving hopes of a Su", suppresses ovary development to produce male- crossed with vinifera to improve propagation. The Jashemski, W. F. (1973). Large vineyard discovered in grape culture where none could survive before. Some ness. A recessive allele, Su', results in reflexing of the ancient Pompeii. Science, N.Y., 180, 821 - 30. Japanese and Chinese cultivars arc oriental species filament and in sterile pollen, to produce functional breeding of rootstocks was the first massive improve- Jelenkovic, G. and Olmo, H. P. (1969). Cytogenetics of introgressed with vinifera. femaleness. In natural populations, males (Su'Sum) ment programme, but the germ plasm of the vinifera Viris. V. Allotetraploids of V. vinifera x V. rotundifoha Wine grapes are seedy, acid and juicy. Among the and females (Su"Sum) occur in equal numbers and scions remained intact. Vitis, 8, 265-79. dessert grapes, scedlessness has evolved in a wide cross-pollination by wind and bees occurs. The domi- Grafting, however, is expensive. Many breeders Lavie, P. (1970). Contribution a fitude caryosystematique range of expression (Stout, 1936). On the one hand are nance relationship of the three alleles is Su" > Su' > therefore set out to breed new vines that would com- des Vitacies. These, Faculte des Sciences de Montpellier, 1, pp. 213. cultivars that have very small, parthcnocarpic berries bine resistance to phylloxera with fruit of good wine Levadotoc, L. (1946). Etude de la fleur et dc la sexualite chez such as the ancient Greek variety Black Corinth, Practically all cultivars of Europe and the New quality, the direct producer. After almost a century, la vigne. dried to produce currants. The Kishmish or Sultanina, Ann. Leek Nat. Agr. Montpellier, 27, pp. 89. World are hermaphrodites (Su e Su* or Su * Sum) and this ideal still remains a dream and we speak now of Neagu, M. M. (1968). Genetique et amelioration de la the most important raisin variety, is said to be steno- self-pollinating. In Middle Asia, however, many arc 'French hybrids'. However, some of the hybrids proved vigne. Rapport general. Off. lit. Vigne ei Yin. Bull., 41, spermocarpic, abortion of the seeds occurring soon female. In a warm, dry climate and with close planting valuable in other ways; for example, in having better 1301 -37. after fertilization. Seedless varieties arise by somatic of mixed cultivars, cross-pollination is effective. As resistance to fungus diseases and greater hardiness., Negi, S. S. and Olmo, H. P. (1971). Conversion and deter- In some areas they were better adapted than ordinary mination of sex in Vitis vinifera (sylvestris). Vitis, 9, Fig. 86.2 Evolution of the grapes, Kris and Muscadinia. vinifera and produced wine of passable quality. They 265-79. Negrul, A. M. (1938). Evolution of cultivated forms of these female cultivars are moved to new areas and form the base of new wine industries in many parts of grapes. C. R. Acad. Sri., U.S.S.R., 18, 585-8. Al usradiuia isolated, they are rapidly selected against because of the world where vinife•ra is not well adapted. The Olmo, H. P. and Baris, C. (1973). Obtention de raisins de 3 spp., North America poor fruitfulness. Female vines are useful in breeding, starting point was a female vine, selection 70 (rupestris 60 ecospecies table apyrines. 0.l.V. mini Symp., Cyprus, 32-11. 2,, = 2.r - eliminating the need for emasculation (Levadoux, x sent by Jaeger from Missouri to Contas- dioccious, interfertile Patel, G. 1. and Olmo, H. P. (1955). Cytogenetics of Vitis. ( wild romarbfaia sot in France in 1882. Contassot distributed open- = 2.v = 38 1946). I. The hybrid V. vinifera x V. rotundifolia. Amer. J. Bot., 'North and Central America, Asia No haploids (n = 19) have been reported in Vitis pollinated seed to Couderc and Seibel who produced 42, 141-59. and the only aneuploids (2n+2) appear as rare abet- the first series of hybrids. The work continues and Stout, A. B. (1936). Seedlessness in grapes. N.Y. Agric. local cvs rants. Autotetraploids were first described in 1929 some of the more recent hybrids have germ plasm Exp. Sta. tech. Bull., 238, pp. 68. USA, seventeenth century ( and have been found to arise spontaneously in most from as many as six American species, but backcrossing cultivars. They are often periclinal chimeras; only to vinifera is still practised to improve quality. For a summary of breeding programmes and accomplish- domestication 400011.c. sterile hybrids two layers are involved in meristem differentiation, so geographical spread 2n 39 that three types of tetraploid chimera have been ments, particularly in Europe, refer to Neagu (1968). Su • mutation 13 ❑ 7 13 I found: 2-4, 4-4 and 4-2. The larger berry size Grapes are outbreeders. Cultivars are highly hetero- attracted attention to the possibility of producing zygous and carry a heavy load of deleterious recessives. improved table grapes but, in general, the auto- Inbreeding depression is severe so that, by the second specialized cvs repeatial auming tctraploids have poor cultural characteristics, being or third generation, sterility usually ensues. The most often hcrm ph., Sa 4 America and Asia own roots, later as scions natural and experimental less fruitful, irregular in berry size and more fragile; successful breeding method is to maintain heterozygo- sity by crossing the best representatives of unrelated (- wine 'table anddried also, the root system is weaker and tetraploids are better grafted on diploid rootstocks. However, a few lines, resorting occasionally to closer mating to acid sweet hybrid cvs tetraploid varieties have long been grown commer- concentrate desirable combinations of characters. seed? less seedy rinilero backcrosscs juicy firm flesh cially in greenhouses where special attention to widely adapted yarthenocarpv., pollination and cultural factors is possible. Alio- 5 Prospects tetraploids ( Jelenkovie and Olmo, 1969) are more Cultivar improvement is increasingly directed toward promising, since the undesirable features of the auto- disease and insect resistance. A high priority is given auto - allo - is tctraploids are not so evident and selection can proceed to virus resistance, since many of the world's oldest little tnc I potential in a wider genetic base. Triploids are highly sterile and most renowned vineyards are seriously menaced but may be useful for vigorous rootstocks, especially by soil-borne infections. Native species must be more if a wide range of resistance to soil pests is desired. thoroughly studied, screened and compared as sources Pentaploids are weak and useless. of resistance. Allopolyploidy as a tool to produce larger Vinifera grapes were propagated from earliest berry size and better cultural features has been times by cuttings or layering and remained relatively neglected in the quest for improved table varieties. free of pests and diseases. However, vines began to We should see the use of native tropical species as a die in French vineyards in 1860 and, in 1868, a root base in greatly extending the zone of commercial grape aphid, Phylloxera, was identified as the cause. This culture. As before, vinifera must be used to introduce insect had been introduced from the USA where it high quality. Hardy clones having short growth lived as a natural symbiont on tolerant native vines. cycles should further extend the range. A beginning Within a few years, thousands of acres were ruined has been made with oriental amurensis but American and, eventually, nearly all the vineyards of Europe species can also be used. Selected female cultivars were in trouble. It was noticed that American hybrids to obtain mass hybridization can be useful. Increasing need for mechanization of harvesting, pruning and To remember the animal and plant classifications, remember the following statement:

"Killer phylloxera creates orders for grapevine suppliers."

Kingdom Phylum Class Order Family Genus Species

Napa Valley College BS/92 OF THE GRAPEVINE

KINGDOM: Plant

DIVISION: Tracheophyta Vascular plants (Phylum) SUB-DIVISION: Spermatophyta (Sub-Phylum) Seed plants CLASS: Angiospermae Flowering, enclosed ovules

SUB - CLASS: Dicotyledonae 2 cotyledons, flower parts 4s or 5s or multiples

ORDER: Rhamnales 7 families, mostly vine-like

FAMILY: Vitaceae Grapevine family, 10 genera GENUS: Vitis About 60 species

SPECIES: vinifera Vitis vinifera - the cultivated, European or Old World grapevine

2_ 0 THE GENUS VITIS

OLD WORLD GRAPE: Vitis vinifera L

AMERICAN VITIS: The following species have contributed to grapevine breeding or are important potential sources of germplasm.

Vitis aestivalis Michaux. Massachusetts and southern New Hampshire to Michigan southward to central Missouri and Georgia.

Vitis berlandieri Planchon. Limestone soils of southwestern Arkansas and through Texas into northeastern Mexico.

Vitis californica Bentham. Along streams in central and northern California and southern Oregon (only of local interest).

Vitis candicans Englemann, Western Arkansas and Louisiana, Oklahoma, central and — eastern Texas, and northern Mexico, mostly on limestone soils.

Vitis champini Planchon. Central and southern Texas, in limy soils.

I/ids cinerea Engelmann. Central states, Louisiana to Wisconsin, on river banks, bottom land, and pond margins.

Vitis cordifolia Michaux. In thickets and along streams in the area from Pennsylvania to eastern Kansas and south to Texas and Florida.

Vitis doaniana Munson. Chiefly northwestern Texas, Oklahoma and New Mexico. (Native habitat about the same as that of V. solonis.)

Vitis girdiana Munson. Along streams in southern California (only of local interest).

Vitis lincecumii (Linsecomii) Buckley. High post-oak lands of southwestern Missouri, northern and eastern Texas, and western Louisiana.

Viti.s. longii. Prince (V. solonis. Hort. Berol). Western Oklahoma, northern Texas, eastern New Mexico, and southeastern Colorado.

Vitis monticola Buckley. Limestone hills of southwestern Texas.

Vitis riparia. Michaux (V. vulpina, Linnaeus). Nova Scotia and New Brunswick to Manitoba, west to the Rock Mountains, south into Texas, and east to Arkansas, Tennessee, and Virginia.

Vitis. rufotomentosa Small. Sandy soils, Florida to Louisiana.

Viti.s. rupestris. Scheele. Sandy stream banks, low hills and mountains, southern Missouri, Illinois, Kentucky, and western Tennessee to southwest Texas. Asian Vitis

Minor importance in cultivation Great interest as source of germplasm for grapevine breeding

About 30 species, examples include:

V. cognetiae

V. thumb ergi

✓ amurensis

✓ armata

Genus Muscadinia (3 Species)

Muscadinia rotundifolia Michaux. (Southern U.S.A.) Muscadine Grape

Muscadinia munsoniana Simpson. (Central and Southern Florida)

Muscadinia popenoei Fennell. (Mexico, rare sp.)

BOTANICAL FAMILY TREE OF THE GRAPEVINE

Species Prominent Cultivars VITIS (Including Hybrids) A Geographic Appreciation aestivalis Delaware, Norton argentif olia arizonica berlandieri baileyana Harm Jan de Blij californica candicans champini cinerea cordifolia doaniana gigas girdiana helleri North illex American indica (cariboea) Species labrusca Concord, Niagara, Catawba lincecumii longii monticola novae•angliae palmata riparia rut otomentosa rupestris shuttleworthii smalliana simpsoni Genus sola VITIS treleasei vulpine Elvira, Clinton

amurensis armata betulifolia coignetiae davidii embergeri Cabernet Sauvignon ficifolia Pinot Noir flexuosa Chardonnay lanata Eurasian Riesling pagnucii Species Zinfandel pedicellata Chenin Blanc pentagona Sauvignon Blanc piasezkii Grenache reticulata SdrniHon romaneti Sylvaner rotordi Nebbiolo thunbergii Trebbiano Pinotage Palomino

North Genus munsoniana American MUSCADINIAE popenoei Species rotundifolia Scuppernong, Eden

BOTANICAL FAMILY TREE OF THE GRAPEVINE

Family Vitaccae (formerly: Arnpelidaccae)

r . 1 1 Genera Vitts Ampelopsu Cissus Parthenocissus 6 other genera ... (94 species) (350 species) (15 species)

Subgenera Euvitis (true grapes) Muscadinia (3 species)

Groups of European (1 species) American Asiatic species (± 20 species) (10-15 species) I • V. rotund:folic: Species Vitus vinifera V. nparia V. rupestris V. berlandieri

Subspecies V. vinifera ssp. sativa V. vinifera (several thousand varieties) ssp. sylvestris Gloire Varieties W. Riesling Furmint Sauvignon bl. du Lot Resseguier de Montpellier (several hundred varieties) Petit Sauvignon Subvarieties none Nemes Furmint Gros Sauvignon

Nomenclature and classification of vitaceae. Clone 239 G.m. Pecs 2 none

Practical Viticulture