Use of Ampelographic Methods in the Identification of Nova

USE OF AMPELOGRAPHIC METHODS IN THE IDENTIFICATION OF NOVA

SCOTIAN GRAPE (VITIS SPP.) CULTIVARS

Laura A. Wiser

Thesis

submitted in partial fulfillment of the

requirements for the Degree of

Bachelor of Science with

Honours in Biology

Acadia University

April, 2011

This thesis by Laura A. Wiser

is accepted in its present form by the

Department of Biology

as satisfying the thesis requirements for the degree of

Bachelor of Science with Honours

Approved by the Thesis Supervisor

______(David Kristie) Date

Approved by the Head of the Department

______(Donald Stewart) Date

Approved by the Honours Committee

______(Sonia Hewitt) Date

I, Laura A. Wiser, grant permission to the University Librarian at Acadia University to reproduce, loan or distribute copies of my thesis in microform, paper or electronic formats on a non-profit basis. I, however, retain the copyright in my thesis.

______Signature of Author

______Date

iii

ACKNOWLEDGEMENTS

There are many people who have helped me make this project possible. First, I would like to thank my supervisor, Dr. David Kristie, for his help and suggestions in conducting my research and in writing my thesis. I would also like to thank Dr. Jonathan

Murray and Kim Strickland at Muir Murray Estate Winery for allowing me to conduct my research in a beautiful work environment over the summer. I would like to thank

NSERC for funding my summer work.

Many people have helped make this project possible by letting me sample plants from their vineyards, and by sharing their knowledge with me. I would like to thank

Walter and Ralf Wuhrer for sharing their expertise in viticulture, and for allowing me to spend many hours in their vineyard sampling plants. I would also like to thank Andrew

Jamieson at Agriculture Canada, Gina Haverstock at Gaspereau Winery, and Jürg Stutz at

Domaine de Grand Pré for allowing me to sample their plants, and for giving me helpful tips along the way.

Finally, I would like to thank the staff at the E.C. Smith Herbarium for allowing me to use space in the herbarium, and for teaching me how to press, dry and properly store my plants.

TABLE OF CONTENTS

LIST OF FIGURES ix

LIST OF TABLES xiii

ABSTRACT xv

1.0 INTRODUCTION 1

1.1 History of Grape Cultivation in Nova Scotia 1

1.2 Factors Involved in Nova Scotian Grape Cultivation 2

1.3 Description of the Vitis Genus 3

1.4 Introduction to Vitis vinifera L. 4

1.5 Introduction to Hybrids 4

1.5.1 French-American Hybrids 5

1.5.2 American Hybrids 7

1.5.3 Other Species and Hybrids 10

1.6 Introduction to Ampelography 10

1.6.1 Introduction to Ampelographic Methods 11

1.6.2 Indument of Plant Parts 12

1.6.3 Colour of Vegetative Plant Parts 13

1.6.4 Leaf Contour and Surface 13

1.6.5 Fruit Characteristics 14

1.6.6. Additional Ampelographic Characteristics 14

1.7 Description of Ampelometric Calculations 15

1.8 Objectives of this Study 17

2.0 MATERIALS AND METHODS 25

2.1 Cultivars and Plants Sampled in this Project 25

2.2 Ampelographic Observations 26

2.3 Number of Leaves Used Per Cultivar 28

2.4 Ampelometric Measurements and Calculations 28

2.5 Ampelographic and Ampelometric Descriptions of Unknown Plants 29

2.6 Graphing and Statistical Analysis 29

3.0 RESULTS 31

3.1 Indument 31

3.1.1 Indument of Growing Tips and Upper and Lower Leaf Surfaces 31

3.1.2 Vein Indument 33

3.1.3 Indument of Petioles and Shoots 33

3.2. Colouration of Hair and Vegetative Plant Parts 34

3.2.1 Hair Colour 34

3.2.2 Colour of Vegetative Plant Parts 34

3.3 Description of Petiolar Sinuses 35

3.4 Dentation 35

3.5 Other Characteristics 35

3.6 Ampelometric Measurements 36

3.7 Description of Unknowns 37

4.0 DISCUSSION 71

4.1 Evaluation of Ampelographic Characteristics 71

4.1.2 Growing Tip and Leaf Indument 71

4.1.3 Indument of Veins, Petioles and Shoots 75

4.1.4 Colour of Hair and Vegetative Plant Parts 76

4.1.5 Petiolar Sinus Description 77

4.1.6 Other Characteristics 78

4.2 Evaluation of Ampelometric Characteristics 79

4.2.1 Vein Length Ratios 79

4.2.2 Sum of Angles 80

4.2.3 Leaf Length to Width Ratios 81

4.2.4 Leaf Size 81

4.2.5 Depth of Superior Lateral Sinuses 81

4.2.6 Depth of Inferior Lateral Sinuses 82

4.3 Comparison to Galet’s Ampelographic Observations 82

4.4 Identification of Unknowns 86

4.4.1 Identification of Unknown #1 86

4.4.2 Identification of Unknown #2 87

4.4.3 Identification of Unknown #3 89

4.4.4 Identification of Unknown #4 89

4.5 Sources of Error and Future Work 90

5.0 CONCLUSIONS 99

6.0 REFERENCES 101

APPENDIX 1 104

APPENDIX 2 106

APPENDIX 3 108

APPENDIX 4 110

vii

APPENDIX 5 112

APPENDIX 6 114

APPENDIX 7 116

APPENDIX 8 118

APPENDIX 9 120

APPENDIX 10 122

APPENDIX 11 123

APPENDIX 12 124

APPENDIX 13 125

APPENDIX 14 126

viii

LIST OF FIGURES

Figure Figure Title Page Number Number

Figures Accompanying Introduction

1 Underside of an upper young NewYork Muscat leaf with 18 felty indument.

2 Underside of a Sovereign Coronation leaf with downy 18 indument.

3 Underside of a mature Riesling leaf with cobwebby 19 indument.

4 Valiant growing tip showing white hair with pink margins. 19

5 Veins on the underside of an upper young leaf showing 20 rough pubescence.

6 Bullate upper surface of a lower young V. amurensis leaf. 21

7 Crimped upper surface of a mature V. amurensis leaf. 21

8 Open growing tip from a De Chaunac plant. 22

9 Globular growing tip from a Sovereign Coronation plant. 22

10 Different tooth shapes present on mature Vitis spp. leaves 23 where (a) is a convex tooth, (b) is concave, (c) is pointed and (d) is hooked.

11 Lyre-shaped petiolar sinus on a mature KW97-8 leaf. 23

12 Underside of a mature Chardonnay leaf with a naked base. 24

13 Main veins, angles and lateral sinuses of a mature KW97-8 24 leaf.

Figures Accompanying Results

14 Changes in indument during development on upper surfaces of 41 leaves of plants with cobwebby growing tips.

15 Changes in indument during development on lower sides of 42 leaves of plants with cobwebby growing tips.

16 Changes in indument during development on upper sides of 43 leaves of plants with downy growing tips.

17 Changes in indument during development on undersides of 44 leaves of plants with downy growing tips.

18 Changes in indument during development on upper sides of 45 leaves of plants with felty growing tips.

19 Changes in indument during development on undersides of 46 leaves of plants with felty growing tips.

20 Node of a Sovereign Coronation plant showing an 52 uncharacterized hair type.

21 Mean L2 to L1 vein length ratios for each cultivar. Error bars 59 indicate one standard deviation away from the mean. A central bolded line represents the mean for the overall population. One and two standard deviations away from the overall mean are also represented with dotted lines. Data presented in this figure corresponds to data in Appendix 1. Raw data is shown in Appendix 10.

22 Mean L3 to L1 vein length ratios for each cultivar. Error bars 60 indicate one standard deviation away from the mean. A central bolded line represents the mean for the overall population. One and two standard deviations away from the overall mean are also represented with dotted lines. Data presented in this figure corresponds to data in Appendix 2. Raw data is shown in Appendix 10.

23 Mean L4 to L1 vein length ratios for each cultivar. Error bars 61 indicate one standard deviation away from the mean. A central bolded line represents the mean for the overall population. One and two standard deviations away from the overall mean are also represented with dotted lines. Data presented in this figure corresponds to data in Appendix 3. Raw data is shown in Appendix 10.

24 Mean sum of angles between L1 and L3 for each cultivar. Error 62 bars indicate one standard deviation away from the mean. A central bolded line represents the mean for the overall population. One and two standard deviations away from the overall mean are also represented with dotted lines. Data presented in this figure corresponds to data in Appendix 4. Raw data is shown in Appendix 11.

25 Mean sum of angles between L1 and L4 for each cultivar. Error 63 bars indicate one standard deviation away from the mean. A central bolded line represents the mean for the overall population. One and two standard deviations away from the overall mean are also represented with dotted lines. Data presented in this figure corresponds to data in Appendix 5. Raw data is shown in Appendix 11.

26 Mean leaf length to width ratios for each cultivar. Error bars 64 indicate one standard deviation away from the mean. A central bolded line represents the mean for the overall population. One and two standard deviations away from the overall mean are also represented with dotted lines. Data presented in this figure corresponds to data in Appendix 6. Raw data is shown in Appendix 12.

27 Mean leaf size for each cultivar. Error bars indicate one standard 65 deviation away from the mean. A central bolded line represents the mean for the overall population. One and two standard deviations away from the overall mean are also represented with dotted lines. Data presented in this figure corresponds to data in Appendix 7. Raw data is shown in Appendix 13.

28 Mean depth of superior lateral sinuses for each cultivar. Error bars 66 indicate one standard deviation away from the mean. A central bolded line represents the mean for the overall population. One and two standard deviations away from the overall mean are also represented with dotted lines. Data presented in this figure corresponds to data in Appendix 8. Raw data is shown in Appendix 14.

29 Mean depth of inferior lateral sinuses for each cultivar. Error bars 67 indicate one standard deviation away from the mean. A central bolded line represents the mean for the overall population. One and two standard deviations away from the overall mean are also represented with dotted lines. Data presented in this figure corresponds to data in Appendix 9. Raw data is shown in Appendix 14.

Figures Accompanying the Discussion

30 Groupings based on the indument of leaves of plants with cobwebby 93 growing tips.

31 Groupings based on the indument of leaves of plants with downy 93 growing tips.

32 Groupings based on the indument of leaves of plants with felty 94 growing tips.

xii

LIST OF TABLES

Table Table Title Page Number Number

Tables Accompanying Results

1 Cultivar groupings based on the type of indument present 40 on growing tips.

2 Groupings based on the indument of veins on 47 undersides of upper young leaves.

3 Groupings based on the indument of veins on 48 undersides of lower young leaves.

4 Cultivar groupings based on the indument of veins on 49 undersides of mature leaves.

5 Cultivar groupings based on indument of petioles. 50

6 Cultivar groupings based on shoot indument. 51

7 Cultivar groupings based on the colour of upper young leaves. 53

8 Cultivar groupings based on colour of shoots and petioles. 54

9 Cultivar groupings based on degree of overlap by petiolar lobes. 55

10 Cultivar groupings based on absence or presence of naked 56 bases on mature leaves.

11 Groupings of cultivars based on tooth size. 57

12 Groupings of cultivars based on tooth shape. 58

13 Indument of upper and lower surfaces and veins of unknown 68 leaves.

14 Ampelographic characteristics of unknown cultivars. 69

xiii

Tables Accompanying Discussion

15 Distinguishing characteristics of certain cultivars. 95

16 An approach to cultivar identification for those cultivars that 96 do not have one specific distinguishing characteristic.

xiv

ABSTRACT

In recent years the evaluation of suitable grape (Vitis spp.) cultivars for the Nova

Scotia grape growing and wine industries has become increasingly important. Despite this, little material exists documenting the morphological characteristics of grape cultivars grown in Nova Scotia. This lack of material could make identifying unknown cultivars in a vineyard problematic. The objectives of this study were to describe grape cultivars based on several ampelographic characteristics proposed by Pierre Galet (1979), and to evaluate these characteristics to determine their usefulness in identification.

Twenty-nine cultivars that are currently grown in Nova Scotia were described based on morphological characteristics such as leaf colour, hair type, tendril placement and tooth size and shape. Leaf measurements, such as vein length ratios, leaf size and sinus depth, were also used to describe cultivars. Results demonstrated which characteristics were useful in identification and which were not. For instance, indument, or hairiness, was useful as it allowed all cultivars to be divided into numerous small groups, and in some cases narrowed down the identities of single cultivars, such as Einset and KW96-1.

Other characteristics, such as tendril placement, were less useful as all sampled cultivars showed an identical pattern. In many cases vein length ratio measurements were also of little use, as these measurements tended to differ little between cultivars. The results of this study provided a preliminary means of cultivar identification and an approach to identification that did not previously exist for Nova Scotian viticulture. However, future work is required to test this proposed method of identification. Additional cultivars and characteristics should also be included in future work.

1.0 INTRODUCTION

1.1 History of Grape Cultivation in Nova Scotia

Nova Scotian wine and table grape cultivation was initially established in the early sixteen-hundreds by French colonists. The first vineyard in Canada was planted in

LaHave, Nova Scotia, by Isaac de Razilly, who came to the area in 1632 to found a

French settlement. Although table grape cultivation persisted throughout much of this historical period, wine grape growing declined after the deportation of the Acadians, as these grapes were primarily cultivated by them. Wine grape growing was not seriously revisited until the mid-twentieth century when Agriculture Canada began testing grape cultivars at its experimental farm in Kentville, Nova Scotia (Murdoch, 1865 ; Naugler and Wright, 2006).

The Kentville experimental farm tested 175 wine and table grape cultivars from

1913 to 1983. Unfortunately, no suitable wine grapes and only one suitable table grape cultivar resulted from these trials (Naugler and Wright, 2006). However, promising cultivars such as New York Muscat, De Chaunac and Maréchal Foch were acquired in the 1960s from the Horticultural Experiment Station in Ontario and were brought to Nova

Scotia. Cultivars were evaluated at various experimental plots established throughout

Nova Scotia. Test plants in all experimental plots did not mature fully. However, it was noted that plants at the Kings County plot matured more fully compared to those at the

Digby and Annapolis County plots (Naugler et al., 2004).

Dr. Roger Dial created the first Nova Scotian farm winery in 1981, which resulted in a renewed interest in wine grape cultivation and additional research at the Agriculture

Canada Kentville Research Station in Kentville, Nova Scotia (Naugler et al., 2004). This

1 increased interest in cultivation and research led to the creation of numerous vineyards throughout Nova Scotia in subsequent years (Naugler and Wright, 2006).

1.2 Factors Involved in Nova Scotian Grape Cultivation

Grape cultivation is generally not possible at latitudes above . Nova Scotia is only slightly below this upper limit, between latitudes of and 8 (Galet, 2000a ;

Murdoch, 1865 p.2). Although Nova Scotia could be considered unfavourable for grape production, one must note that some of the principal wine producing regions of France, such as Alsace, Bordeaux, Burgundy and Champagne, are located between these latitudes

(Williams et al., 2009 p. 78) However, Nova Scotia generally has warmer summers and colder winters than these areas. A wide range of climate and soil characteristics also exists throughout the province, meaning that some areas are suitable for grape production whereas others are not. Regions that are particularly favourable for grape production include Bear River Valley, Annapolis Valley, LaHave River Valley, Gaspereau Valley,

Halifax County, Avon River Valley, Malagash Peninsula and Antigonish (Naugler et al.,

2004).

The suitability of a region for grape production is largely based on the number of heat units an area experiences. In viticulture, heat units are the sum of average daily temperatures above C from April 1st to October 31st (Mullins et al., 1992). Nine- hundred heat units are generally required to ripen early Vitis vinifera cultivars.

Greenwood, for example, has an average of 950 heat units, and Kentville has an average of 964. Other considerations for planting grapevines in Nova Scotia are that vine leaves can only withstand frosts of - C, and most cultivars require 150 days at temperatures above - C to fully mature. Greenwood, for example, has an average of only 123 frost

2 free days and Kentville has an average of only 134. However, it should be noted that frost free days in these instances were considered to be those days at or below C not at or below - C. One would therefore expect there to be slightly more than 123 days in

Greenwood and slightly more than 134 days in Kentville where temperatures are above -

C. A third consideration is that most hybrid and V. vinifera cultivars exhibit differing susceptibilities to cold temperatures, and often have specific minimum temperatures that they can be exposed to before experiencing injury or death. Cold hardiness of a cultivar is therefore an important consideration for vineyards in Nova Scotia. (Naugler and Wright,

2006).

1.3 Description of the Vitis Genus

The genus Vitis is comprised of 60 to 70 members. Members are perennials that possess a climbing habit made possible by flexible shoots and pressure-sensitive tendrils.

These tendrils are always located opposite leaves (Keller,2010 ; Mullins et al.,1992). The emergence of tendrils indicates that the grapevine has acquired a climbing habit, but not necessarily a state of maturity. Leaves are palmate, where all five main veins arise from a single point. The five main veins make up the five lobes of the leaf (Mullins et al., 1992).

Members of the genus Vitis are best suited for areas with warm, dry summers and cold wet winters, and grow primarily in the temperate regions of the Northern

Hemisphere (Mullins et al., 1992 ; Keller,2010). Vitis vinifera L. is the only species that originated from Eurasia. This species was distributed throughout much of the world by humans (Mullins et al., 1992).

1.4 Introduction to Vitis vinifera L.

Vitis vinifera L., also known as the European grape, is a temperate-climate species that cannot survive particularly cold winter weather and requires hot summers to produce fully-matured berries. Cultivars of Vitis vinifera are generally considered to be the foundation of wine and table grape cultivation (Mullins et al., 1992). A cultivar is comprised of a number of identical plants that have been selected for particular advantageous traits. When propagated vegetatively, the plants of a cultivar retain those traits (Keller, 2010). Conversely, varieties are comprised of a number of plants with identical traits, where these traits have appeared without human selection. Unlike cultivars, most varieties propagate themselves sexually (Gordon, 1997 p.4 ).

Chardonnay and Riesling are two out of a small number of Vitis vinifera cultivars grown in Nova Scotia (Naugler and Wright, 2006). Both are used to produce white wine

(Galet, 2000b). Riesling is suitable for cultivation in Nova Scotia as it is one of the most cold-tolerant Vitis vinifera cultivars. However, one of the challenges of growing Riesling in Nova Scotia is that the cultivar is late-ripening, where full maturation is not generally achieved until late October. Chardonnay ripens in early to late October and is less cold- tolerant than Riesling. Chardonnay’s cold-tolerance is often considered to be inadequate for use in Nova Scotian vineyards. Despite this, the cultivar has been grown at a number of vineyards in the province (Naugler and Wright, 2006).

1.5 Introduction to Hybrids.

Grapevine hybrids are produced by crossing different cultivars or species, usually to obtain progeny with desirable characteristics. Crosses are normally written with the maternal parent followed by the paternal parent (Keller, 2010). Unfortunately, the

4 parentage of most cultivars is unknown (Galet, 2000a). It is also customary to assign new hybrids a letter and number code rather than a proper name. The letter represents the site where the hybrid was developed or the name of the breeder, and the first few numbers represent the year in which the hybrid was created. Numbers after this are assigned by the breeder. Once the hybrid is deemed to be marketable it is given a proper name (Naugler et al., 2004).

Members of the genus Vitis experienced a high frequency of naturally-occurring hybrids throughout their evolutionary history, and therefore achieved a considerable amount of morphological diversity. The need to produce grapevine hybrids did not arise until grapevine diseases spread throughout Europe in the late nineteen hundreds (Mullins et al., 1992).

European colonists originally brought Vitis vinifera to the Americas in the 17th century (Galet, 2000a; Mullins et al.,1992). However, these grapes did not flourish as they were susceptible to pests present in the New World, such as phylloxera,

Daktulosphaira vitifoliae Fitch, an insect that feeds on the roots and leaves of grapevines.

These grapevines could also not withstand cold temperatures. In the nineteenth century, colonists brought American species back to Europe, and in doing so accidentally introduced American pests, such as powdery mildew in 1845 and phylloxera in 1868.

This resulted in the devastation of the European grape industry (Galet, 2000a).

In order to create cultivars that were both resistant to phylloxera and could produce suitable wine, breeders crossed V. vinifera cultivars, which were susceptible to phylloxera, with phylloxera tolerant or resistant American species. The resulting

5 grapevines are known as French-American hybrids (Pool et al., 1979; Mullins et al.,

1992).

1.5.1 French-American Hybrids

French-American hybrids resulted from crosses of American species other than

Vitis labrusca L. (Pool et al., 1979). American species in these crosses include Vitis riparia Michaux, Vitis rupestris Scheele and Vitis aestivalis Michaux. Vitis riparia has the earliest budbreak of the three species and is commonly used as a parent because of its early maturation, its phylloxera-resistant roots and its resistance to cold temperatures.

Like Vitis riparia, Vitis rupestris is also commonly used in crosses because of its early maturation and its phylloxera-resistant roots. Vitis aestivalis is used in crosses because of its large berries compared to many other American species. However, unlike V. rupestris and V. riparia, V. aestivalis is not resistant to phylloxera (Galet, 1988).

The French-American hybrids examined in this project were Maréchal Foch,

Léon Millot, Luci Kuhlmann, Triomphe d’Alsace, Baco Noir, Precoce de Colmar, Seyval

Blanc, Vidal Blanc and De Chaunac. Maréchal Foch (Kuhlmann 188-2), Léon Millot

(Kuhlmann 194-2) and Luci Kuhlmann (149-1 Kuhlmann) were produced from a cross between 101-13 Mgt and Goldriesling, making them Vitis riparia - Vitis rupestris - Vitis vinifera hybrids (Galet, 1979; Galet, 1988; Galet, 2000b). All three cultivars are suitable for cultivation in Nova Scotia as they are cold hardy. Maréchal Foch and Léon Millot are resistant to most diseases, whereas Luci Kuhlmann is susceptible to bunch rot and powdery mildew (Woods, 2006; Naugler and Wright 6). Triomphe d’Alsace is another Kuhlmann hybrid, however its parentage is unknown (Naugler and Wright,

2006). Information was also difficult to obtain for Precoce de Colmar, or Kuhlmann 277-

2. According to the Vitis International Variety Catalogue (http://www.vivc.de), this hybrid was produced by crossing Knipperlé with Millardet et Grasset 101- 14.

Baco Noir, also known as Baco 1 or 24-23 Baco, resulted from a cross between the Vitis vinifera cultivar Folle Blanche and Vitis riparia (Galet, 1979; Galet, 2000b). It may not be as suitable for growth in Nova Scotia compared to other red hybrids, as it ripens later and is less resistant to colder temperatures. Seyval Blanc is also difficult to grow in Nova Scotia. Of all the hybrids in this study, it is considered to be the least resistant to cold and is generally more susceptible to disease (Naugler and Wright, 2006).

This cultivar, also known as Seyve-Villard 5.276, resulted from a cross between S.5656 and S.4986 (Galet, 1979).

Vidal Blanc, or Vidal 256, resulted from a cross between Ugni Blanc and S. 4986

(Galet, 1979). This hybrid has only moderate cold- tolerance as well as late fruit ripening

(Naugler and Wright, 2006). The origin of De Chaunac, or Seibel 9549, is largely unknown. It is an early-ripening and cold-hardy hybrid that likely resulted from a cross between Seibel 5163 and Seibel 793 (Galet, 1979; Galet, 2000b; Weaver, 1976 p. 97).

1.5.2 American Hybrids

Unlike French-American hybrids, nearly all American hybrids involve crosses where Vitis labrusca L. is a parent (Pool et al., 1979). V. labrusca has a distinct appearance. It has growing tips and leaf undersides that are covered in dense hair. Hair is usually white, except on the growing tips where hair is normally pink. Leaves are often described as being shield-shaped, or cuneiform (Galet, 1979). Because of their cold hardiness, Vitis labrusca hybrids are particularly important in the wine industry of Nova

Scotia (Fisher and Jamieson, 2000; Galet, 1998). The American V. labrusca wine grape

7 hybrids used in this project were L’Acadie Blanc New York Muscat Cayuga and

Frontenac. The American V. labrusca table grape hybrids used in this project were

Canadice, Einset, Reliance, Swenson Red and Valiant.

L’Acadie Blanc is considered by many to be the most renowned grape cultivar grown in Nova Scotia (Naugler and Wright, 2006). This cultivar was created at the

Horticultural Research Institute of Ontario by crossing Cascade, or Seibel 13053, with

Seyve-Villard 14-287, and was originally given the code V53261. It was deemed to be unsuitable for cultivation in Ontario but was found to be promising in certain areas of

Nova Scotia (Woods, 2006; Fisher and Jamieson, 2000). This hybrid is especially appropriate for cultivation in Nova cotia because it is disease resistant and can withstand temperatures as low as - C. It also grows relatively well in shorter growing seasons (Fisher and Jamieson, 2000; Naugler and Wright, 2006).

Other prominent American hybrids are New York Muscat and Cayuga, both of which were created at the Geneva Experiment Station at Cornell University, New York.

New York Muscat is moderately disease resistant and can withstand temperatures as low as - 6 C. Cayuga is also cold and disease resistant (Rombough, 2002; Naugler and

Wright, 2006; Woods, 2006). Another American wine grape used in the project,

Frontenac, resulted from a cross between Vitis riparia and Landot 4511. This hybrid was developed at the University of Minnesota (Rombough, 2002).

Canadice and Einset, commonly used to produce table grapes, were bred at the

New York Experiment Station at Cornell. Canadice was produced from a cross between

Bath and Himrod whereas Einset was produced from a cross between Fredonia and

Canner. Einset is therefore a Vitis labrusca – Vitis vinifera cross. Like Einset, Reliance is

8 also a Vitis labrusca – Vitis vinifera cross and resulted from a cross of Ontario with

Suffolk Red. Developed in Arkansas, this grape is suitable for Nova Scotian cultivation as it is disease-resistant and tolerant to cold temperatures. Swenson Red and Valiant are also both cold resistant cultivars. Swenson Red was produced from a cross between

Mn78 and 11.803 and Valiant from a cross between a Vitis riparia cultivar and Fredonia

(Galet, 2000b; Rombough, 2002).

Several hybrids produced at the Kentville Agriculture Canada Research Station were also used in this study. The research station began its grape breeding program in

1953, but quickly suspended it. The program resumed in 1983, and in the following year more than 2300 seedlings were planted from 26 different crosses. KW87-1, which resulted from a cross between V.53035 and Castel 19637, was one of the hybrids that showed potential. It was crossed with Seyval Blanc to produce 46 seedlings which were planted at the Agriculture Canada Kentville Research Station in 1989. Out of the hybrids produced from these crossings, KW94-1 to KW94-5 were the most promising. A promising cultivar is one that shows potential for profitability in wine or table grape industries, and one that is suitable for Nova Scotian cultivation. Other promising hybrids were developed in 1991 and 1992 and were selected in 1995, 1996 and 1997. These cultivars are currently being assessed to determine if they are suitable for Nova Scotian wine and table grape growing industries (Naugler et al., 2004). The Kentville hybrids used in this project were KW96-1, which resulted from a cross between KW87-1 and

Ortega , KW96-2, which resulted from a cross between St. Pepin and Siegerrebe, KW97-

4, which resulted from a cross between St. Pepin and Ortega, KW97-8, which resulted from a cross between KW87-2 and Ortega, and KW97-23, which resulted from a cross

9 between St. Pepin and L’Acadie (A. Jamieson, personal communication, November 10,

2010).

1.5.3. Other Species and Hybrids

Cab-Foch, S-4-29 and Sovereign Coronation were also used in this project, but no published material relating to these cultivars could be obtained. According to P. Troop

(personal communication, 06 March 2011), a winemaker in British Columbia, Cab-Foch was bred in Switzerland by Valentin Blattner in the late 1980s or early 1990s. Cab-Foch is believed to be a cross between Cabernet Sauvignon and Maréchal Foch. According to

P. Bowen (personal communication, 04 March 2011), at the Pacific Agri-Food Research

Centre in Summerland B.C., where S-4-29 was likely produced, records were not maintained after breeding programs ended. No information could therefore be found for this cultivar. Sovereign Coronation was also produced at the Summerland Research

Centre, and according to the Vitis International Variety Catalogue (http://www.vivc.de), this cultivar was obtained by crossing Lady Patricia with Himrod in 1966.

V. amurensis was the only Asian species used in this project. This species may be of interest in breeding programs aimed at producing cold-hardy hybrids as it can withstand temperatures as low as - C. However, it may not be suitable in areas where winter temperatures fluctuate. It also has little resistance to American pests (Galet, 2000a;

Rombough, 2002).

1.6 Introduction to Ampelography

Ampelography is used to describe, identify and classify different species and cultivars of the Vitis genus (Keller, 2010). Before the eighteenth century, ampelography was primarily used in monasteries, as wine was of great importance in Christian rituals

(Mullins et al., 1992). At this time descriptions were based only on the aptitudes of grapevines. Plant parts were not used as they were thought to be too variable for use in identification. This posed problems when, during the later half of the nineteenth century, many grapevine diseases were introduced into Europe from America. Numerous

European vineyards were destroyed and needed to be replanted with cultivars that were resistance to these diseases. This compelled people to study these disease-resistant species and cultivars, however, there was no established means of cultivar identification.

It was not until 1944 when Pierre Galet and H. Agnel first began creating the first key to identify certain cultivars, which ultimately led Galet to develop ampelography (Galet,

1979).

1.6.1 Introduction to Ampelographic Methods

Galet (1998) deemed different plant parts to be of varying importance in identification and description. The shoot tip is considered to be the most important plant part in identification, and should therefore be described first, followed by young and mature leaves. Young leaves are considered to be the upper three leaves that are clearly separated from the shoot tip. The lower young leaves are those leaves that are still immature, but are below the top three. Shoots and petioles are considered after leaves.

Inflorescences and flowers, grape clusters, berries and seeds are of less importance and should be described last. The morphological characteristics of each plant part are considered independently and there is a separate morphological classification for each part (Galet, 1998).

1.6.2 Indument of Plant Parts

Indument, or hairiness, is a primary ampelographic characteristic. Indument of the growing tip is crucial as it determines the indument of young and mature leaves (Mullins et al., 1992). Hairs on grapevines can be categorized as being either woolly or pubescent.

Woolly hairs are present in different densities. Indument is considered to be felty when hair is so densely distributed that one is unable to determine the colour of the underlying plant material (Figure 1). If one can easily distinguish the colour of the underlying plant material, the hair should be classified as downy or cobwebby. Indument is considered to be downy when hair is densely distributed but one can easily determine the colour of the underlying plant material (Figure 2). Indument is considered to be cobwebby when hair is not densely distributed (Figure 3). The colour of woolly hairs can be described as white, tawny (yellowish-brown), rust (reddish-brown), pink or white with pink margins (Figure

4; Galet, 1998).

If hairs are short and erect they are considered pubescent, and pubescent hair can be velvety or rough. Plant parts that have slightly hooked short hairs are displaying rough pubescence (Figure 5), whereas those with densely distributed straight hairs are displaying velvety pubescence. A hairless plant part is said to be glabrous (Galet, 1998).

A grapevine leaf has a somewhat constant number of hairs throughout development.

As leaf size increases the hairs therefore decrease in density. For instance, a plant with a felty growing tip will likely show high densities of hair on the youngest leaves, but only downy or cobwebby indument on lower young and mature leaves (Galet, 1979). The indument of the growing tip, because it directly relates to the indument of the young and

12 mature leaves, is considered to be an important ampelographic characteristic (Galet,

1998).

1.6.3 Colour of Vegetative Plant Parts

Colour of plant parts can be an important ampelographic characteristic, however can also be quite subjective and can vary depending on nutritional status. Upper young leaves are generally described as being pale green, yellowish-green, red, copper, or green with copper areas. Leaves below this are generally different shades of green. Younger leaves that are pale green gradually become a deeper shade of green as they mature

(Galet, 1998). Shoots and petioles are generally green, however in many cultivars and species shoots and petioles are red on surfaces exposed to the sun. The colour of the growing tip is only visible if it is glabrous or lightly woolly. The colour of the growing tip is generally the same as that of upper young leaves (Galet, 1979).

1.6.4 Leaf Contour and Surface

Leaf contour is the general 3D shape of the leaf. Leaves are described as flat when surfaces are somewhat even and folded when leaves are creased along the midvein.

Leaves are concave or involute when the leaf edges turn upward and convex or revolute when the leaf edges turn downward. Leaves are considered contorted when they have an overall irregular shape. The appearance of the leaf surface is a secondary ampelographic characteristic. Bullate leaves have small bumps that are only a few millimetres in diameter (Figure 6), and crimped leaves have bumps that are at least one centimetre in diameter (Figure 7; Galet, 1979).

1.6.5 Fruit Characteristics

Fruit description is only of secondary importance in ampelography as fruit can vary considerably based on growing conditions. The length and weight of fruit clusters can be noted. Fruit clusters less than 6cm long would be considered very small, whereas fruit clusters more than 24cm long would be considered very large. In terms of weight, a cluster weighing less than 50 grams would be very small, whereas a cluster weighing more than 1000 grams would be considered very large (Galet, 1979).

The shape of fruit clusters can be described as cylindrical, where primary lateral branches are of equal length. Cone-shaped or conical clusters have lateral branches that become smaller as proximity to the tip decreases. Shouldered clusters have large lateral branches near the peduncle and winged clusters have lateral branches nearest the peduncle that are large and separated. Cluster density, which describes the amount of space between berries, can also be considered. This is based on the size of the berries, the length of the pedicels and the structure of the lateral branches (Galet, 1979).

1.6.6. Additional Ampelographic Characteristics

Tendril placement can be considered when describing a grapevine. Continuous shoots are those that have tendrils located at every node, except the first three or four at the base of the shoot. Subcontinuous shoots are those that have at least three successive nodes with tendrils. Intermittent or discontinuous shoots never have more than 2 tendrils in succession (Galet, 1979).

The shape of the growing tip, which is located at the shoot apex, should also be noted. It can be open (Figure 8), where leaves are separate from the tip, or globular, where leaves are not separate from the growing tip (Figure 9; Galet, 1979).

Dentation on mature leaves should be described based on size and shape. Size is determined by dividing tooth length by width. If this ratio is equal to or less than 0.25, the tooth is considered to be almost non-existent. If the length to width ratio is between 0.26 and 0.5 the tooth is considered to be wide. An average tooth is one that has a length to width ratio of 0.51 to 0.75, a narrow tooth is one that has a ratio between 0.76 and 1, and a very narrow tooth is one that is longer than it is wide. Tooth shape should be noted as convex, concave, pointed or hooked (Figure 10).

1.7 Description of Ampelometric Calculations

Ampelometric calculations are used to better demonstrate the shape of a leaf.

These calculations alone cannot be used to identify a cultivar. However, calculations are a valuable tool when used in conjunction with other characteristics. The length of each main vein and their ratios, the leaf size, leaf length to width ratio, and the angles between main veins are normally calculated (Galet 1998).

Members of the genus Vitis have 5 main veins (Figure 13; Keller, 2010). The midvein (L1) runs from the petiolar junction up the center of the leaf. The superior lateral vein (L2) is the vein closest to the midvein, followed by the inferior lateral vein (L3). The petiolar vein (L4) is not directly connected to the petiolar sinus, but is instead connected to the inferior lateral vein (Figure 13). The lengths of all main veins should be measured.

The length of a vein is obtained by measuring the distance between the petiolar junction and the point where it meets its corresponding tooth (Galet, 1998). One can then determine the vein length ratios by dividing the lengths of L2, L3, and L4 by the length of L1. Angles between L1 and L2, L2 and L3 and L3 and L4 can also be determined.

Galet (1979) assigned these values the symbols α β and γ respectively (Figure 13).

Values of high importance are the sum of α and β as well as the sum of all three measurements. Leaf size can vary considerably, however one can still obtain a general idea of size by multiplying leaf width by length. The length can also be divided by the width to give a length to width ratio (Galet, 1998).

Ampelometric calculations can be used to determine a leaf’s shape. There are basic leaf shapes. Cordiform leaves are heart-shaped, where L1 is the longest vein.

Cuneiform leaves are shield-shaped, where the sum of the angles is rarely more than

. Like cordiform leaves, cuneiform leaves are always longer than they are wide.

Truncate leaves are similar to cuneiform leaves, but have shorter L3 veins and larger angles. The length to width ratio, in this case, normally varies from 0.9 to 1.1. Orbicular leaves have a circular appearance, where the petiolar junction is at the center of the circle and all main vein angles are roughly equal to 6 . Reniform leaves are kidney-shaped, where the sum of the angles is never more than and the leaf is wider than it is long.

Between these five basic leaf shapes, there are also intermediate combinations (Galet

1979).

Leaf lobes and sinuses are also important leaf descriptors. Each main vein has a corresponding lobe with a sinus between it. The sinus between L1 and L2 is the superior lateral sinus (SS), the sinus between L2 and L3 is the inferior lateral sinus (SI), and the sinus that lies between the two petiolar veins is the petiolar sinus (Figure 13). One should take into consideration the depths of the lateral sinuses. This is done by calculating the ratio between the size of the lobe and the size of its corresponding vein

(Galet, 1998).

One can also note the shapes of the lobes. Lobes can overlap completely, covering the sinus, partially, where the sinus is still partly visible, or not at all. Lobes can also be described as lyre-shaped (Figure 11), U or V-shaped. Mature leaves can also have naked bases, where there is no plant material below the area where L3 and L4 join

(Figure 12; Galet, 1979).

1.8 Objectives of this Study

The objective of this study was to describe cultivars grown in Nova Scotia based on ampelographic characteristics and ampelometric measurements. The morphological characteristics of most Nova Scotian cultivars have not been documented in full, making identification difficult. Characteristics and measurements obtained from each cultivar will be evaluated to determine their usefulness in identification. Useful characteristics and measurements will be used to create a preliminary means of identification.

Figure 1. Underside of an upper young NewYork

Muscat leaf with felty indument.

Figure 2. Underside of a Sovereign Coronation leaf

with downy indument.

Figure 3. Underside of a mature Riesling leaf with

cobwebby indument.

Figure 4. Valiant growing tip showing white hair

with pink margins.

Figure 5. Veins on the underside of an upper young

leaf showing rough pubescence.

Figure 6. Bullate upper surface of a lower young

V. amurensis leaf.

Figure 7. Crimped upper surface of a mature V. amurensis

leaf.

Figure 8. Open growing tip from a

De Chaunac plant.

Figure 9. Globular growing tip from a

Sovereign Coronation plant.

Figure 10. Different tooth shapes present on mature Vitis spp. leaves where (a) is a convex tooth, (b) is concave, (c) is pointed and (d) is hooked.

Figure 11. Lyre-shaped petiolar sinus on a

mature KW97-8 leaf.

Figure 12. Underside of a mature Chardonnay

leaf with a naked base.

Figure 13. Main veins, angles and lateral sinuses of a mature

KW97-8 leaf.

2.0 MATERIALS AND METHODS

2.1 Cultivars and Plants Sampled in this Project

Observations were based on a number of representative plants from each of the twenty-nine cultivars used in this study. The healthiest available plants were chosen from various vineyards in the Annapolis Valley region of Nova Scotia. Only mature plants, those that were able to produce fruit, were used. Each plant was labelled with a ribbon and assigned a number. Baco Noir, Chardonnay, Precoce de Colmar and Triomphe d’Alsace had three representative plants each all of which came from Muir Murray

Estate Winery. Cayuga, Einset Seedless, Reliant, Sovereign Coronation and S-4-29 had three representative plants each all of which were located at Walter Wuhrer’s North

Mountain vineyard. Cab-Foch also had three representative plants, which came from

Domaine de Grand Pré. De Chaunac, Lucie Kuhlmann, Riesling and Vidal Blanc had three representative plants each, all of which came from the Gaspereau Winery.

Canadice and KW96- had three plants each two of which came from Walter Wuhrer’s vineyard and one of which came from the Atlantic Food and Horticulture Research

Centre in Kentville, Nova Scotia. Cultivars with only one representative plant were

Frontenac, KW96-1, KW97-4, KW97-8, KW97-23, Swenson Red, Valiant and V. amurensis, which came from the Atlantic Food and Horticulture Research Centre.

Cultivars with four representative plants were L’Acadie Blanc Léon Millot Maréchal

Foch, New York Muscat and Seyval Blanc. Each of these plants had 3 representatives from Muir Murray Estate Winery and one representative plant from the Atlantic Food and Horticulture Research Centre.

Three shoot samples were obtained from each plant and pressed in a herbarium press provided by the E.C. Smith Herbarium. At least two mature leaf samples per plant were also pressed. These samples were stored at the E.C. Smith Herbarium.

All characteristics of interest from each cultivar were photographed. This included upper and lower sides of upper young, lower young and mature leaves. Photos were also taken of shoots, petioles and growing tips.

2.2 Ampelographic Observations

Choice of characteristics to include in this project was based on A Practical

Ampelography: Grapevine Identification (Galet, 1979). Observations were made from mid-June to mid-September. Morphological characteristics were based on observations from shoot and leaf samples that were removed from plants to be pressed and dried, in addition to observations from at least 3 other shoots or leaves per plant. Each plant was described separately, starting with the growing tip. In this study, the leaves that had just separated from the growing tip were described in conjunction with the growing tip. The indument of the growing tip was noted as being either glabrous, cobwebby, downy, felty or having rough or velvety pubescence. Hair colour was noted as being either white or white with pink margins. There were instances where tawny-coloured hair was observed on various plant parts. These instances, however, were only observed part way through the season and appeared on different plant parts at seemingly random times. This color was therefore not considered in results. Growing tips were noted as being either open or globular.

The upper young leaves, which were considered to be the three upper leaves that were clearly separated from the growing tip, were described as being either glabrous,

26 cobwebby, downy, felty or as having rough or velvety pubescence. Plant parts that had only a few sparse hairs were considered to be nearly glabrous. A plant part was considered to be glabrous to nearly glabrous, or glabrous - nearly glabrous, when some of the plant parts were hairless and others had very few hairs. The indument of upper and lower sides of leaves was considered independently. The indument of the veins on the undersides of leaves was also noted. This was repeated for lower young and mature leaves. Indument that transitioned from one type to another within one developmental section, for instance mature leaves that were cobwebby but gradually become glabrous, were noted as being cobwebby to glabrous, or cobwebby-glabrous. Leaf surface was also noted as being either bullate or crimped. Upper young, lower young and mature leaves were considered separately.

Indument of petioles and shoots was also noted as being glabrous, cobwebby, downy, felty, or as having rough or velvety pubescence. Shoots and petioles of the entire plant were described together. The colour of shoots and petioles was also noted as being either red, green, or a mixture of both colours.

Mature leaves were used to obtain descriptions of the petiolar sinus and dentation.

The petiolar sinus of mature leaves was described as being lyre, U, or V-shaped, and fully, partially or non-overlapping. The shape of teeth was noted as being either convex, concave, hooked or pointed. Tooth size was noted as being barely-existing, wide, average, narrow or very narrow. One tooth per leaf was measured by dividing its length by width. Other teeth on the same leaf were then compared to this measurement to determine a general tooth size.

Tendril placement was described as being continuous, subcontinuous or intermittent/ discontinuous. At least 3 shoots were observed per plant to determine this pattern.

2.3 Number of Leaves Used Per Cultivar

Ampelometric calculations were based on a number of mature leaves from each representative plant. Each leaf was given a number. Three leaves were sampled per each representative Cab-Foch, Cayuga, Einset Seedless, Reliant, Sovereign Coronation, S-4-

29, Baco Noir, Chardonnay, Precoce de Colmar, Triomphe d’Alsace De Chaunac Luci

Kuhlmann, Riesling, Vidal Blanc, Canadice, and KW96-2 plant. This resulted in there being 9 leaves for each of these cultivars. Cultivars with four representative plants,

L’Acadie Blanc Léon Millot Maréchal Foch New York Muscat and eyval Blanc also had 3 leaves per plant, resulting in there being 12 leaves for each cultivar. Ten leaves were taken from each cultivar with only 1 representative sampled plant. These were

Frontenac, KW96-1, KW97-4, KW97-8, KW97-23, Swenson Red, Valiant and V. amurensis.

2.4 Ampelometric Measurements and Calculations

Measurements were taken from live or dried leaf specimens. All measurements were taken using a simple ruler and protractor. The length of all main veins, L1 through

L4, was measured. Ratios were determined by dividing the lengths of L2, L3, and L4 by the length of L1. Angles between L1 and L2, L2 and L3, and L3 and L4 were measured.

The sum of the angle between L1 and L2 was determined in addition to the sum of all angles. The length to width ratio of each leaf was determined by dividing the length of L1 by the length of the section of leaf that appeared to have the largest width. These length

28 and width values were then multiplied together to obtain an overall approximate leaf size.

The depth of the superior and inferior lateral sinuses was also determined. To measure the depth of the superior lateral sinus, the distance between the petiolar sinus and the most indented area of the leaf between L1 and L2 was obtained and then divided by the length of the L2 vein. To measure the depth of the inferior lateral sinus the distance between the petiolar sinus and the most indented area of the leaf between L2 and L3 was obtained and then divided by the length of the L3 vein.

2.5 Ampelographic and Ampelometric Descriptions of Unknown Plants

Four unknown plants were chosen from Muir Murray Estate Winery. Each unknown was assigned a number . Unknowns 1, 2 and 3 were found amongst rows of

Seyval Blanc. Unknown 4 was found amongst a row of New York Muscat. Ten leaves were sampled for Unknowns 1, 3 and 4. Eight leaves were sampled for Unknown 2 as there were very few mature leaves available on this plant. Procedures from sections 2.2,

2.4 and 2.6 were applied to unknowns.

2.6 Graphing and Statistical Analysis

Measurements were analyzed using GraphPad Prism (Version 5.0a, Graphpad

Software Inc. San Diego, CA). Mean values and standard deviations were based on measurements from L’Acadie Blanc Léon Millot, Maréchal Foch, Seyval Blanc and

New York Muscat leaves, 10 Frontenac, KW96-1, KW96-2, KW97-4, KW97-8, KW97-

23, Swenson Red, Valiant, V. amurensis, Unknown 1, 2 and 3 leaves. Mean values and standard deviations from Unknown 2 were based on measurements from 8 leaves, and the means and standard deviations of all other cultivars were calculated based on 9 leaves per cultivar. The mean values from each cultivar were graphed, along with error bars

29 representing one standard deviation away from the mean. One-way ANOVA tests with

Tukey’s post tests were used to compare data between each cultivar. A confidence interval of 99.9% was used due to the large number of comparisons. At higher alpha values some comparisons could be significantly different by random chance.

Additionally, more significant differences would emerge at lower confidence intervals, making patterns in data more difficult to discern. This data was then transferred into matrices, displayed in Appendices 1-9, which displayed all possible comparisons between cultivars. In these matrices, significantly different comparisons were noted as “ ” and not significantly different comparisons were noted as “ ”.

3.0 RESULTS

3.1 Indument

The indument of the growing tip, upper and lower sides of upper young, lower young and mature leaves was noted, followed by the indument of veins, petioles and shoots. Plant parts were noted as being glabrous, cobwebby, downy, felty, or as having rough pubescence. Velvety pubescence was not observed on any plant part.

3.1.1 Indument of Growing Tips and Upper and Lower Leaf Surfaces.

Plants with glabrous, cobwebby, downy and felty growing tips were observed.

Pubescence was not observed on growing tips. Grape cultivars were divided into four primary groupings based on the indument present on growing tips (Table 1).

Most sampled cultivars had felty growing tips. Only two cultivars, De Chaunac and Frontenac, had glabrous growing tips. The growing tips of other cultivars were either cobwebby or downy. Cab-Foch, Chardonnay, Riesling and Vidal Blanc fit into two different categories. Cab-Foch could be considered cobwebby or downy. Chardonnay,

Riesling and Vidal Blanc had growing tips that could be considered either downy or felty.

Growing tip indument is important as it directly relates to the indument of all other grapevine leaves. As leaves expand, they generally have a set number of hairs, as maturing leaves do not produce new hairs. Hair density will therefore decrease as leaves expand during maturation (Galet, 1979). One can therefore further divide the four primary groupings in Table 1 based on changes in indument during maturation.

As would be expected, the two plants with glabrous growing tips from Table 1,

De Chaunac and Frontenac, had glabrous upper young leaves, lower young leaves and mature leaves on both upper and lower surfaces.

Cultivars that had cobwebby growing tips had glabrous to cobwebby upper sides of upper young leaves. Hair became less dense as maturation progressed, however, this decrease in density occurred at differing rates depending on cultivar. Figure 14 shows the changes in indument that occurred during development on upper sides of leaves from cultivars that had cobwebby growing tips. Leaf undersides from plants that had cobwebby growing tips were glabrous or nearly glabrous and, as expected, remained so throughout maturation. Figure 15 shows the changes in indument during development on lower sides of leaves from cultivars with cobwebby growing tips.

Cultivars that had downy growing tips were generally cobwebby on upper sides of upper young leaves and had hair that became less dense as leaves matured. Chardonnay was the only cultivar that had downy upper sides of upper young leaves. However, these leaves quickly became cobwebby, resulting in there being both downy and cobwebby indument on lower young leaves (Figure 16). Leaf undersides were either downy or glabrous at the youngest stages of growth (Figure 17). Those with downy or downy- cobwebby young leaves remained either downy, cobwebby or nearly glabrous throughout development.

The upper sides of upper young leaves from plants that had felty growing tips were felty, downy or cobwebby (Figure 18). Hair density decreased as leaves matured.

The undersides of upper young leaves were felty, downy or cobwebby. There were far less instances of cobwebby hair on undersides of leaves compared to upper sides. Most leaf undersides were felty and became downy as leaves matured (Figure 19).

3.1.2 Vein Indument

Indument of veins on leaf undersides appeared to be unrelated to the indument of the growing tip and was therefore considered independently from other leaf surfaces.

Upper young, lower young and mature leaves were considered separately.

Most cultivars had upper young leaves with veins that had more than one hair type (Table 2). Veins were usually pubescent with one type of woolly hair. Frontenac was the only cultivar with rough pubescence that did not also have woolly hair. L’Acadie

Blanc, De Chaunac and KW97-23 were the only cultivars that were glabrous or nearly glabrous.

A similar pattern was observed on lower young leaves, except that there was a decrease in woolly hair density, resulting in there being additional leaves with cobwebby indument and fewer leaves with downy indument (Table 3). Canadice was the only cultivar with lower young leaves that had veins with downy hair.

A similar pattern was again observed on mature leaves (Table 4). There were, however, fewer leaves with woolly indument. Many leaves had veins with only pubescent hair.

3.1.3 Indument of Petioles and Shoots

Petiole indument was also considered. Most cultivars exhibited more than one type of indument (Table 5). Whenever more than one woolly hair type was present, denser hair types were observed on younger plant parts and less dense hair types were observed on more mature plant parts. Shoot indument was similar to petiole indument, however, pubescent hair appeared on petioles of fewer cultivars compared to shoots

(Table 6). Cultivars with rough pubescence on shoots were KW96-2, Léon Millot, New

York Muscat and S-4-29.

In addition to the previously-mentioned hair types, Sovereign Coronation exhibited an unnamed indument that was not present on any other cultivar (Figure 20).

3.2 Colouration of Hair and Vegetative Plant Parts

3.2.1 Hair Colour

When present, hair was white. The only exceptions were pink margins on growing tips as well as on leaves that had just separated from the growing tips of certain cultivars.

Cultivars with prominent pink margins were Luci Kuhlmann, Canadice, Valiant, V. amurensis and Triomphe d’Alsace. Reliance Riesling eyval Blanc Maréchal Foch

Precoce de Colmar, Chardonnay, Cab-Foch, Sovereign Coronation, KW96-1 and KW96-

2 had less prominent pink margins that were not always present or noticeable.

3.2.2 Colour of Vegetative Plant Parts

Upper young leaves were green, red, or green with red areas (Table 7). Four cultivars, De Chaunac, Einset, KW97-4 and KW97-8, had entirely red upper young leaves. Another four cultivars L’Acadie Blanc Frontenac wenson Red and V. amurensis, had both green and red areas on upper young leaves. All other upper young leaves were predominantly green.

Petioles and shoots were green, red or green with varying degrees of red (Table

8). Red areas were generally only present on plant surfaces that were exposed to sunlight.

There were five cultivars L’Acadie Blanc De Chaunac Frontenac, Riesling and V. amurensis, that were entirely red. Other cultivars were green, or more often green with varying degrees of red coloration.

3.3 Description of Petiolar Sinuses

All cultivars had lyre-shaped petiolar sinuses. Some Chardonnay, Precoce de

Colmar and Baco Noir leaves also had U-shaped petiolar sinuses. The degree to which petiolar lobes overlapped was also used to create cultivar groupings (Table 9). Vidal

Blanc and KW97-23 were the only cultivars with sinuses that completely overlapped.

Other cultivars generally showed no overlap on some leaves and partial overlap on others.

The absence or presence of naked bases on mature leaves was also noted (Table

10). Most cultivars did not have naked bases. Cab-Foch, Frontenac, KW97-4, Luci

Kuhlmann, Maréchal Foch and Triomphe d’Alsace showed naked bases on some mature leaves but not others. Chardonnay was the only cultivar where naked bases were consistently present on every observed leaf.

3.4 Dentation

Most cultivars had leaves with teeth of various sizes (Table 11). Most cultivars also had leaves with more than one tooth shape (Table 12). Only 4 cultivars, Léon

Millot, Luci Kuhlmann, Sovereign Coronation and Precoce de Colmar, had hooked teeth.

However, these cultivars had other tooth shapes in addition to the hooked teeth.

3.5 Other Characteristics

Upper young leaves were generally slightly bullate to bullate. Exceptions were

Canadice, Cayuga, De Chaunac, Einset Seedless, Frontenac, KW96-1, KW96-2, and S-4-

29, which had smooth surfaces. Most lower young leaves had smooth surfaces, with the only exception being V. amurensis, which had bullate to crimped leaves. Most mature leaves were also smooth. Exceptions were L’Acadie Cab-Foch, KW97-4, KW97-8 and

KW97-23, Luci Kuhlmann, Riesling, Triomphe d’Alsace Vidal Blanc and V. amuresis, which were slightly crimped. Most leaves that were bullate or crimped had only very small bumps. It was therefore difficult to differentiate between smooth and bullate or crimped leaves. The only cultivar that was distinctly crimped and bullate was V. amurensis.

All cultivars had a discontinuous tendril pattern, where there were never more than two nodes with tendrils in succession. Nearly all cultivars also had an open growing tip. Exceptions that had distinctly globular tips were L’Acadie Blanc overeign

Coronation and KW97-23.

3.6 Ampelometric Measurements

The majority of mean L2 to L1 vein length ratios were similar amongst cultivars.

Mean L2 to L1 vein length ratios for V. amurensis and Vidal Blanc were significantly smaller than the ratios of many other cultivars (Appendix 1; Figure 21). Most mean L3 to

L1 vein length ratios were also similar amongst cultivars. The mean L3 to L1 ratios of

V.amurensis and Vidal Blanc were significantly smaller than the ratios of several other cultivars (Appendix 2; Figure 22). Cab Foch also appeared to have a larger mean L3 to

L1 ratio compared to many other cultivars. Most mean L4 to L1 vein length ratios were also similar amongst cultivars. The mean values from V.amurensis and Vidal Blanc were again significantly smaller than several other cultivars (Appendix 3; Figure 23). The mean value from Riesling was significantly larger than that of many other cultivars, such as Baco Noir, De Chaunac, Luci Kuhlmann and New York Muscat.

The mean sum of angles between L1 and L3 generally did not differ significantly between cultivars. From Appendix 4 and Figure 24, V. amurensis and Frontenac were

36 shown to have significantly smaller L1 to L3 angles compared to many other cultivars and Vidal Blanc had significantly larger angles compared to many other cultivars. The mean sum of angles between L1 and L4 showed slightly more variation. Figure 25 and

Appendix 5 showed that Einset and Vidal Blanc had L1 to L4 angles that were significantly larger compared to many other cultivars. Frontenac, Luci Kuhlmann and V. amurensis had smaller angles compared to many other cultivars.

Mean leaf length to width ratios generally did not differ significantly between cultivars (Figure 26; Appendix 6). V. amurensis, however, had a significantly larger mean ratio compared to all but 2 cultivars, not including unknowns. These two cultivars were Baco Noir and Precoce de Colmar.

Mean sizes varied significantly between cultivars (Figure 27; Appendix 7).

Sovereign Coronation was significantly larger than all other cultivars in this study.

Canadice was also significantly larger than all but 3 cultivars, which were Einset,

Sovereign Coronation and Reliance.

A number of cultivars had mean superior lateral sinus depths that were significantly different from most other cultivars (Figure 28; Appendix 8). Canadice and

KW97-8 had sinuses that were significantly deeper compared to most other cultivars. In addition, Appendix 8 shows that the values from New York Muscat and KW97-23 were significantly different from those obtained from most other cultivars. The mean depths of inferior lateral sinuses generally did not differ significantly between cultivars. The only exceptions were KW96-1 and KW97-8, which, according to Figure 29 and Appendix 9, had significantly deeper inferior lateral sinuses compared to most other cultivars.

3.7 Description of Unknowns

Unknown 2 and 4 had cobwebby growing tips whereas Unknown 1 and 3 had felty growing tips. Unknown 1 and 3 showed a similar pattern of felty and downy young surfaces which gradually became less dense as maturation progressed. Veins for both were also downy and gradually became cobwebby. Unknown 2 and 4 had upper young leaves with upper surfaces that were cobwebby and gradually became glabrous. The undersides of these leaves were glabrous with pubescent veins. Table 13 shows all indument patterns on leaf surfaces and veins. All other ampelographic observations are shown in Table 14.

Unknowns 1 and 3 had mean L2 to L1 vein length ratios that were significantly different from the vein length ratios of many other cultivars (Figure 21; Appendix 1).

The mean vein length ratio of Unknown 2 did not differ significantly from the mean values of any other cultivar. Mean L3 to L1 vein length ratios for Unknowns 1, 2 and

3were significantly different from several other cultivars (Figure 22; Appendix 2).

Unknown 4 showed no significant difference compared to any other cultivars in this case.

All unknowns showed no or almost no significant difference in mean L4 to L1 vein length ratios compared to other cultivars (Figure 23; Appendix 3).

The mean sum of angles between L1 and L3 from Unknown 4 was significantly smaller than the sum of angles from many other cultivars (Figure 24; Appendix 4).

Angles obtained from Unknown 1 and 3 were significantly larger than several other cultivars. The mean sum of angles between L1 and L3 for Unknown 2 was not significantly different from the means of any other cultivars. Similar patterns were shown in Figure 25 and Appendix 5.

The mean leaf length to width ratios obtained for each unknown cultivar were comparable to the means of most other cultivars (Figure 26; Appendix 6). Unknown size was also comparable to most other cultivars (Figure 7; Appendix 7). Lobes of unknown cultivars were not significantly deeper than many other cultivars (Figure 8; Figure 9;

Appendix 8; Appendix 9).

Table 1. Cultivar groupings based on the type of indument present on growing tips.

Glabrous Cobwebby Downy Felty (Hairless) (Few hairs) (Dense hairs, but (Hair so dense that it is underlying plant colour difficult to determine can be determined.) underlying plant colour.)

De Chaunac L’Acadie Blanc Baco Noir Canadice Frontenac **Cab Foch **Cab Foch Cayuga KW97-23 *Chardonnay *Chardonnay Léon Millot KW97-8 Einset Seedless Maréchal Foch Luci Kuhlmann KW96-1 Precoce de Colmar *Riesling KW96-2 Seyval Blanc S-4-29 KW97-4 Swenson Red Triomphe d’Alsace New York Muscat Valiant *Vidal Blanc Reliant *Riesling Sovereign Coronation *Vidal Blanc Vitis amurensis *Cultivars that fit into both downy and felty groups. **Cultivars that fit into both cobwebby and downy groups.

s of plants with cobwebby growing tips. growing plants of s with cobwebby

of leave of

during development on upper surfaces upper on development during