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This is to certify that the

dissertation entitled

Self-Incompatibility in Sour Cherry (Prunus cerasus L.)
and Inbreeding and Multivariate Relationships Among
Almond (Prunus dulcis (Miller) D.A. Webb) Cultivars

 

 

presented by

ALI LAN SARI

has been accepted towards fulfillment
of the requirements for

Ph.D degree in Horticulture

 

 

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SELF-INCOMPATIBILITY
IN SOUR CHERRY (Prunus cerasus L.)
and
INBREEDING AND MULTIVARIATE RELATIONSHIPS AMONG
ALMOND (Prunus dulcis (Miller) D.A. Webb) CULTIVARS
BY

Ali Lansari

A DISSERTATION

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

DOCTOR OF PHILOSOPHY

Department of Horticulture

1993

 

ABSTRACT
SELF-INCOMPATIBILITY
IN SOUR CHERRY (Prunus cerasus L.)
and
INBREEDING AND MULTIVARIATE RELATIONSHIPS AMONG
ALMOND (Prunus dulcis (Miller) D.A. Webb) CULTIVARS
BY

ALI LANSARI

Self-incompatibility was investigated in sour cherry
(Prunus cerasus L.) by examining pollen growth in the pistil
using ultraviolet (UV) fluorescence microscopy following self-
and cross—pollination. The sour cherry cultivars
'Tschernokorka' and ‘Crisana' exhibit pollen tube inhibition
in the style characteristic of gametophytic self—
incompatibility. ‘Meteor' and ‘Montmorency' appear to be
partially self-incompatible with few self-pollen tubes
reaching the ovary. Pollen growth rate is different according
to the male parent and to the receptive pistil.

Pedigrees of 124 almond, Prunus dulcis (Miller) D.A Webb,
cultivars from U.S., Russia, Israel, France, and Spain were
used to calculate: l) the inbreeding coefficients of these
cultivars, 2) the genetic coancestries among these cultivars,
and 3) the genetic contribution of founding clones to these
cultivars. The recurrent use of 4 selections as parents in the
U.S. breeding programs has resulted in a mean inbreeding
coefficient (F) within the U.S. germplasm collection of 0.022.

In France, a single cultivar, 'Ferralise', has an inbreeding

 

value of F = 0.250, while cultivars of other almond producing
countries are noninbrafl (F=0). Due to the use of common
parents, the American, Russian, and Israeli cultivars share
coancestry, while coancestries also exist between French and
Spanish almond germplasm. Cultivars of known parentage,
including those released through breeding programs, in the
U.S., Russia, Israel, France, and Spain trace back
respectively to 9, 8, 3, 4, and 3 founding clones. Almond
breeding programs might, in the future, narrow the almond

germplasm genetic base thereby limiting genetic gain.

Principal component analysis (PCA) was used to quantify
morphological variation among eighty one selected Moroccan
clones and introduced cultivars. Moroccan selections tended
to be characterized by small leaves in comparison to foreign
cultivars. Variability for nut and kernel traits was
identified, and along with several clones which have very good
nut and kernel characteristics. There is, however, some
evidence that the fruiting potential of Moroccan selections
remains limited, even though some of them had a large number
of spurs. No evidence was found of separate ecotypes existing

in the southern Moroccan almond populations.

DEDICATION

To my father and mother who always gave me love and
‘support, and whom I always cherished and loved.

To my father who taught me a lot about life.

To my wife, Fouzia, who has always been beside me, and who
always provided me with the love and support I needed
throughout the years of our marriage. She always left
everything behind to help and encourage me to complete this
work.

To my cherished children, Kawtar and Soufiane, who always
brighten my dark moments with their smiles and laughs. I love
them very much.

To my brothers, Aziz and Mohammed, and my sisters, Houria,
Sabah, Zoulikha, Bouchra, and Soumia, for their understanding
and love.

To all my friends.

iv

 

ACKNOWLEDGMENTS

I wish to express my sincere gratitude to my major
professor, Dr. Amy F. Iezzoni, for her advice, support and
understanding throughout the duration of my graduate program
in Michigan. I am most grateful to her for her trust in my
ability to complete the present research in both Michigan,
USA, and Morocco.

I would like to express my sincere appreciation to the
members of my Guidance Committee, Drs. J. Whallon, F. Dennis,
J. Hancock, and J. Kelly.

Deep thanks to Dr. A. Cameron for taking time to read my
thesis and for serving on my examining Committee.

I am thankful to Dr. Dale E. Kester, from the University of
California, Davis, for providing me with advice, ideas, and
support that I needed to conduct this research.

Thanks to Drs. Ch. Grasselly, R. Socias I Company, and J.

Luby for their critical reading of the manuscript.

 

I . ,;:‘,-,:-s'—”!f(' ‘-

TABLE OF CONTENTS

Page
LIST OF TABLES.... .................................. Vii
LIST OF FIGURES ..................................... viii
LIST OF APPENDICES .................................. ix
INTRODUCTION ................................... ..... 1

CHAPTER I. A PRELIMINARY ANALYSIS OF SELF—
INCOMPATIBILITY IN SOUR CHERRY (Prunus cerasus L.).. 3

Summary ............................................. 4
Introduction ........................................ 4
Materials and methods ............................... 5
Results ............................................. 8
Conclusions ......................................... 13
Literature cited .................................... 15

CHAPTER II. INBREEDING, COANCESTRY, AND FOUNDING
CLONES OF ALMONDS OF CALIFORNIA, MEDITERRANEAN

SHORES, AND RUSSIA ................................. 16
Summary ............................................. 17
Introduction ........ . ............................... 17
Materials and methods ............................... 22
Results .......................................... ... 24
Conclusions ......................................... 38
Literature cited .................................... 42

CHAPTER III. MORPHOLOGICAL VARIATION WITHIN
COLLECTIONS OF MOROCCAN ALMOND CLONES AND

MEDITERRANEAN AND AMERICAN CULTIVARS ......... . ..... 55
Summary ............................................ 56
Introduction .................. . ......... . ........... 56
Materials and methods ......................... ..... 59
Results ........ . .............. ...... ........ ....... 65
Conclusions........ ......... ........ ......... ...... 79
Literature Cited .............. .. ....... . .. ........ 82
GENERAL CONCLUSIONS ............... ... ......... ..... 97

vi

 

 

LIST OF TABLES

Table Page
Chapter I:
1. Self-incompatibility (-) and self-compatibility
among 6 cherry cultivars, based upon controlled
crosses ............................................. 9
Chapter II:
1. Country of origin, parentage, and inbreeding
coefficients for 10 almond cultivars ................ 25
2. The percent genetic contribution of founding clones
over all the cultivars in their respective groups
outlined in the pedigrees in Appendices 2-6.... ..... 27
3. Coefficients of coancestries for U.S. almond
cultivars ........................................... 30
4. Coefficients of coancestries for Russian almond
cultivars used in breeding programs ................. 31
5. Coefficients of coancestries for French almond
cultivars used in breeding programs .............. ... 33
6. Coefficients of coancestries for Spanish almond
cultivars used in breeding programs ............... 34
7. Coefficients of coancestries for Israeli almond
cultivars used in breeding programs ................ 36
7. Mean coancestry coefficients among almond cultivars
from 5 almond producing countries.................. 37
Chapter III:
1. Origin and location of cultivars and clones ........ 61
2. Morphological characters and ratios employed in the
analyses .............................. ......... 64
3. Eigenvectors of the 23 principal components axes
from PCA analyses of Stations 1, 2, and 3 genotypes
represented in Figures 2, 3, and 4 ....... ...... 66
4. Means of representative characters highly loading
on PC1, PC2, and PC3 axes for Figure 2 (station #1) 69
5. Means of representative characters highly loading
on PC1, PC2, and PC3 axes for Figure 3 (station #2) 73
6. Means of representative characters highly loading
on PC1, PC2, and PC3 axes for Figure 4 (station #3) 77

vii

 

LIST OF FIGURES

Figure

Chapter I:

1.

Pollen tube growth in sour cherry pistils 72 hours
after pollination : (top) 'Crisana' self-
pollinated, (middle) 'Meteor' self-pollinated, and
(bottom) 'Crisana' x 'Meteor'.The pistils were
purposely curved before the photographs were
taken ............................................ .
Mean of the percentage of style length of (A)
'Tschernokorka', (B) 'Crisana', (C) ‘Meteor', and
(D) 'Montmorency', penetrated by pollen tubes 24,
48, and 72 hours after pollination. Values
represent the mean of 10 styles per cross .........

Chapter III:

1.
2.

3.

4.

Survey areas map of southern Morocco... ...... .....
Position of PC scores of introduced cultivars and
Moroccan selections. Station #1 ...................
Position of PC scores of introduced cultivars and

Moroccan selections. Station #2 ...................
Position of PC scores of introduced cultivars and
Moroccan selections. Station #3 ...................

viii

Page

10

11

6O

68

72

76

 

LIST OF APPENDICES

Appendix Page

Chapter II:

1.

2.
3.
4.
5.
6.

Parentage and inbreeding coefficients of almond
cultivars grown in the U. 5., Russia, Israel, France,

Spain, and Italy .................................. 46
Pedigrees of U. S. almond cultivars ................. 50
Pedigrees of Russian almond cultivars .............. 51
Pedigrees of French almond cultivars ............... 52
Pedigrees of Spanish almond cultivars ........ . ..... 53
Pedigrees of Israeli almond cultivars .............. 54

Chapter III:

10.

11.

12.

13.

Means of characters measured highly loading on PC1

for Figure 2 (Station #1) .......................... 84
Means of characters measured highly loading on PC2

for Figure 2 (Station #1) .......................... 85
Means of characters measured highly loading on PC3

for Figure 2 (Station #1) .......................... 86
Means of characters measured highly loading on PC1

for Figure 3 (Station #1) .......................... 87
Means of characters measured highly loading on PC2

for Figure 3 (Station #1) .......................... 88
Means of characters measured highly loading on PC3

for Figure 3 (Station #1)............. ............ 89
Means of characters measured highly loading on PC1

for Figure 4 (Station #1) .......................... 90
Means of characters measured highly loading on PC2

for Figure 4 (Station #1) .......................... 91
Means of characters measured highly loading on PC3

for Figure 4 (Station #1) .......................... 92
Eigenvalues of the correlation matrix for the

seven first PC's at Station #1 .................... 93
Eigenvalues of the correlation matrix for the

seven first PC's at Station #2 ..................... 94
Eigenvalues of the correlation matrix for the

seven first PC's at Station #3... ............. ..... 95
Almond nuts and kernels of Moroccan selections ..... 96

ix

 

INTRODUCTION

Self-incompatibility ensures outcrossing which promotes
heterozygosity, permitting a greater capacity to adapt to
different environments. This may result in a wider species
range. Self-compatibility can develop through evolution,
either by mutation of self-incompatibility alleles to self-
compatible ones, or by interspecific hybridization of self-
incompatible and self- compatible species, giving rise to
self-compatible genotypes within a species or to a new self—
compatible species. Prunus species are either self-compatible
or self-incompatible. The tetraploid sour cherry (Prunus
cerasus L.) is thought to have resulted from hybridization
between sweet cherry (2; gyigm L.), which is a self—
incompatible diploid, and the tetraploid ground cherry (£1
fruticosa Pall.). Sour cherry is known to be self-compatible,
but cultivars with varying degrees of self-incompatibility
have been reported. The objective of the work described in
chapter 1 was to characterize self-incompatibility in sour
cherry. This was achieved by examining pollen tube growth in
the pistil using ultra violet fluorescence microscopy, in four
sour cherry cultivars and several hybrids of self-compatible
sour cherry cultivars.

Under certain environmental conditions and in the presence
of natural barriers, gene flow can be restricted and species

'can be isolated, giving rise to identifiable ecotypes. Under

 

human and natural selection forces in different countries,
almond ecotypes have evolved and developed characteristic
traits (i.e., self-compatibility in almond (Prunus dulcis
(Miller) D.A. Webb) Puglia ecotypes in Italy, soft shell types
in California). Fruit breeders select superior genotypes of
almond from available ecotypes, and use them as commercial
cultivars and as gene sources for cultivar improvement.
Extensive use of common parents in breeding programs reduces
genetic diversity and increases inbreeding and coancestry
relationships among released cultivars. However, in Morocco,
almond, which is an outcrossing Prunus species, is mostly
propagated by seeds and is grown under various environmental
conditions. These conditions increase genetic diversity and
may result in the development of characterized ecotypes.

In Chapter 2, the level of inbreeding and coancestry
relationships among almond cultivars from different almond
producing countries was determined. In Chapter 3,
morphological variation among Moroccan selections and
introduced cultivars was studied using multivariate

statistics.

 

CHAPTER I

A PRELIMINARY ANALYSIS OF SELF-INCOMPATIBILITY

IN SOUR CHERRY (Prunus cerasus L.)

SUMMARY:

Self-incompatibility was investigated in sour cherry
(Prunus cerasus L.) by examining pollen tube growth in the
pistil using ultraviolet (UV) fluorescence microscopy
following self— and cross-pollination. The sour cherry
cultivars 'Tschernokorka' and ‘Crisana' exhibit pollen tube
inhibition in the style characteristic of gametophytic
self—incompatibility. ‘Meteor' and ‘Montmorency' appear to be
partially self—incompatible since some but not many
self-pollen tubes reach the ovary. Pollen germination rates
were different according to the pollen source and the

receptive pistil.

INTRODUCTION:

Self—incompatibility in Prunus is widespread. Most
commercial almond cultivars (2. dulcis Miller) (Socias I
Company et al., 1976; Crossa—Raynaud and Grasselly, 1985) and

sweet cherry cultivars (P.

avium L.) (Crane and Lawrence,

 

1929; Crane and Brown, 1937; Way, 1967) exhibit gametophytic

self-incompatibility and cross-incompatible groups have been

identified. In plums (2. domestica), the expression of
self-incompatibility is more complex. Plums can be
self-compatible, self-incompatible, or partially
self-compatible, and only a few examples of

 

cross-incompatibility have been reported (Crane and Lawrence,
1929; Suranyi, 1978).

Although the tetraploid sour cherry is generally
considered self-compatible, self-incompatible sour cherry
cultivars do exist (Lech and Tylus, 1983; Redalen, 1984a,
1984b). These self-incompatible cultivars generally represent
old landrace cultivars which are being replaced in commercial
production by self-compatible types. Besides the two
classifications of self-compatible and self—incompatible,
Redalen (1984b) describes a partially self—incompatible class
in sour cherry, which he defined as having 1.5% to 15% fruit
set.

The sour cherry industry in the United States is a
monoculture of the cultivar ‘Montmorency'. Beginning in 1983,
sour cherry germplasm, including some self—incompatible
cultivars, was imported into the United States from Europe for
use in sour cherry breeding. The objectives of this study were
to evaluate pollen tube growth in sour cherry cultivars
previously reported to be self-compatible or
self-incompatible. Pollen tube growth in the pistil was

examined using ultraviolet (UV) fluorescence microscopy.

MATERIALS AND METHODS:

Plant material: A diallel cross was made among four sour

cherry cultivars: ‘Tschernokorka', ‘Crisana', ‘Meteor', and

I , A,,~~1-..u-r

‘Montmorency'. ‘Tschernokorka' and ‘Crisana' are reportedly
self-incompatible (Redalen, 1984a, 1984b). Two sweet cherry
cultivars, ‘Emperor Francis' and ‘Schmidt' were also included
for comparison.

All the plant material tested was planted at the
Clarksville Horticultural Experimental Station, Clarksville,

Michigan, U.S.A.

Pollination and evaluation: Pollen was collected from cut
branches forced indoors at room temperature. Pollen viability
was tested by the method of Werner and Chang (1981) using 3(4—
5-dimethylthiazolyl-2)2,5-diphenyl tetrazolium bromide (MTT)
(Sigma, St. Louis, MO.) as the pollen staining agent, because
of its high correlation with pollen germination. A drop of a
solution of 100 mg of MTT in 5 ml of 10% sucrose solution was
placed on the microscope slide, the pollen dusted onto the
surface, and a coverslip added. Counts were made one hour
later for the development of a satisfactory red color. Pollen
grains that stained red or dark red were considered as viable,
pollen staining light red were considered of reduced
viability, and colorless pollen grains were non viable.

Flowers on branches in the field selected for crossing were
emasculated at the balloon stage, all other flowers being
removed. Emasculated flowers were hand pollinated when
receptive. 'Schmidt' and 'Emperor Francis' were pollinated.May

13, 1989, 'Montmorency', 'Crisana', and 'Tschernokorka' were

 

pollinated May 14, and 'Meteor' was pollinated May 16. The
minimum and maximum temperatures for May 13 through May 19
were 9-15°C, 10-20°C, 10-21°C, 10-25°C, 11-29°C, 14-27°C, and
17-22°C, respectively. Ten pollinated pistils per cross were
collected 24, 48, and 72 hours after pollination and fixed in
Farmer's fixative solution (glacial acetic acidzethanol, 1:2,
v/v).

The pistils were prepared for fluorescent microscopy
with a modification of Martin's technique (1959). They were
thoroughly washed under running tap water and autoclaved for
10-minutes in 1% sodium sulfite (Merck, Cherry Hill, N.J.)
solution to soften the tissues. After removal of the
epidermis, the pistils were soaked in 0.1% aniline blue
(Sigma, St. Louis, MO.) solution at pH 8.5 for 10 to 15
minutes. Pollen tubes were observed by UV fluorescent light
microscopy using an Olympus BH-2 microscope, with A 10 PL of
0.25 aperture, and A40 PL of 0.65 aperture objectives. The
exciter filters used were UGL for ultraviolet excitation, and
BP49OB for blue excitation. Aniline blue in alkaline solution
will fluoresce under UV light when it is complexed with
polysaccharide which is present in callose plugs (Stone et
al., 1984).

Pollen-pistil incompatibility’ was defined as no pollen
tubes reaching the ovule on all pistils. observed, while
compatibility was considered to exist when at least one

observed pistil had pollen tubes reaching the ovule.

 

 

Pollen tube growth was determined by the distance pollen
tubes had penetrated into the upper 1/4, upper 1/3, lower 1/3,
lower 1/4 of’ the style, or' reached. the ovule. Data is
represented as the mean percentage of total style length

travelled by pollen tubes of 10 pistils per cross.
RESULTS:

All cultivars tested were cross compatible (Table 1).

‘Tschernokorka' and ‘Crisana' sour cherries and the sweet
cherry' cultivars *were -self—incompatible; 1K) pollen. tubes
reached the ovule in any of the 10 pistils observed. ‘Meteor'
and ‘Montmorency' were scored; in these two groups, at least
50% of the pistils had pollen tubes reaching the ovule (Table
1). However, not all the pistils observed had pollen tubes
that reached the ovule 72 hours after pollination (Fig. 2C and
D). Self-incompatibility in ‘Tschernokorka' and ‘Crisana' was
characterized by inhibition of pollen tube growth in the
stylar tissue (Fig. 1). Self-pollen of ‘Crisana' and
‘Tschernokorka' initially grew as fast as foreign sour cherry
pollen; however, pollen tube growth was reduced 48 hours after
pollination.and stopped.approximately half way down the styles
72 hours after pollination (Fig. 2A and B). In the
incompatible sour cherry pollinations, pollen tube branching,
bursting or growth reversal were observed. The swelling of

pollen tube tips, which is usually associated with

 

rm. ;.' "'2' z , ' ;.' '

 

Table 1. Self-incompatibility (-) and self-compatibility (+)
among six cherry cultivars based upon controlled

 

 

 

crosses.
Pollen
Pistil EF Sch Tsch Cris Met Mont
Emperor Francis - + + + + +
SChIllidt + - + + + +
Tschernokorka + + - + + +
Crisana + + + - + +
Meteor + + + + + +
Montmorency + + + + + +

 

(-) indicates that pollen tube growth in all pistils observed was arrested
in the stylar tissues.

(+) indicates that more than 50% of the pistils observed had pollen tubes
reaching the ovule.

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'Montmorency' , covered by pollen tubes 24, 48, and

(A)' Tschernokorka', (B) 'Crisana', (C) lMeteor', and
72 hours after pollination.

(D)

2. Mean of the percentage of style length of

Fig.

Values represent the mean of 10 styles.

9 24 h. a 48 h, [J 72 h after pollination.

 

 

 

 

 

 

 

  

 

 

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gametophytic incompatibility, was observed in the sweet cherry
cultivars but not in the self-incompatible sour cherry
cultivars. 'Crisana' pollen grew the best on 'Tschernokorka'
pistils (Fig.2), while all cross pollen types grew succesfully

on 'Crisana' pistils (Fig. 2C).

‘Montmorency' and ‘Meteor' were self—compatible with
self-pollen successfully reaching the ovule. However, for
‘Montmorency', self—pollen grew more slowly than foreign
pollen. Effectively, the mean percentage of total style length
travelled by pollen tubes 72 hours after pollination was 80%,
since not all the pistils observed had pollen tubes that
reached the ovule (Fig. ZD). In the ‘Meteor' styles, self-
pollen and pollen of 'Crisana' and 'Montmorency' generally
grew more slowly, with mean values of 80%, 65%, and 60%
respectively, as compared to pollen of the two sweet cherry
cultivars and 'Tschernokorka'sour cherry (Fig. 2C), and as
compared to the styles of the other sour cherry cultivars
(Fig. 2). Additionally, for both ‘Montmorency' and ‘Meteor',
very few self-pollen tubes reached the last one third of the
style (Fig. 1), many of them being inhibited along the
transmitting tissue jJ: a pattern more variable than that
exhibited for out-cross pollinations (Fig. 1C). This could not
be attributed to pollen quality, since the ‘Montmorency' and
‘Meteor' pollen grew successfully in the styles of other

cultivars. It was noted that sweet cherry pollen generally

12

 

 

grew faster than sour cherry pollen in sour cherry styles

(Fig. 2A-D).

CONCLUSION:

Self-incompatibility in sour cherry is characterized by
inhibition of pollen tube growth in the style suggesting
gametophytic self-incompatibility as reported for other
self—incompatible Prunus species. In the incompatible
pollinations, the pollen tubes initially penetrated into the
style and grew normally; however, growth stopped when the
pollen tubes were approximately half way down the style. Our
results are contrary to those of Lech and Tylus (1983) who
observed no pollen tube growth beyond the stigma for the sour
cherry cultivar ‘Koroser', presumably synonymous to the
Romanian cultivar ‘Crisana'.

Redalen (1984b) classified ‘Montmorency' and ‘Meteor' as
partially self-incompatible based on low fruit set following
self-pollinations. ‘Montmorency' and ‘Meteor' did exhibit
inhibition of a large percentage of self—pollen in the style.
This observation could represent the partial
self-incompatibility described by Redalen (1984a, 1984b).

Pollen growth was different according to the donor parent
and to the female parent. In general, 'Emperor Francis' and
'Schmidt' sweet cherry pollen was rapid in growth on sour

cherry cultivars. Sour cherry pollen had the slowest growth on

13

 

 

'Meteor'. The ploidy level at the gametic stage could affect
pollen tube growth rate between pollen of the diploid sweet
cherry and pollen of the tetraploid sour cherry on sour
cherry pistils. Moreover, partial and complete self-
incompatibility on sour cherry seems to be more complex to
understand based on the presence of a fertility restorating
allele (or alleles) from one of the believed sour cherry
parents, the tetraploid Prunus fruticosa, and considering that

sour cherry is an allotetraploid.

14

 

Literature Cited

Crossa-Raynaud, P. and C. Grasselly. 1985. Existance de
groupes d'intersterilité chez l'amandier. Options
Mediterraneenes 85:43-45.

Crane, M.B. and W.J.C. Lawrence. 1929. Genetical and
cytological aspects of incompatibility and sterility in
cultivated fruits. J. Pom. Hort. Sci. 7:276-301.

Crane, M.B. and. A.G. Brown. 1937. Incompatibility and
sterility in the sweet cherry, Prunus avium L. J. Pom.
Hort. Sci. 15: 86—116.

 

Lech, W. and K. Tylus. 1983. Pollination, fertilization, and
fruit set of some sour cherry varieties. Acta Hort.

139:33-39.

Martin, F.W. 1959. Staining and observing pollen tubes in
the styles by means of fluorescence. Stain Technol.
34:125-128.

Redalen, G. 1984a. Cross pollination of five sour cherry
cultivars. Acta Hort. 149:71-76.

Redalen, G. 1984b. Fertility in sour cherries.

Gartenbauwissenschaft 49(5/6):212-217.

Socias I Company, R., D.E. Kester and M.V. Bradley. 1976.
Effects of temperature and genotype on pollen tube growth
in some self—incompatible and self-incompatible almond
cultivars. J. Amer. Soc. Hort. Sci. 101:490—493.

Stone, B. A., N. A. Evans, I. Bonig, and A. E. Clarke. 1984.
The application of Sirafluor, a chemically defined
fluorochrome from Aniline blue for the histochemical
detection of callose. Protoplasma 122: 191- 195.

Suranyi, D. 1978. A new method to determine self-fertility in
plum varieties. Acta Hort. 74: 155-162.

Way, R.D. 1967. Pollen incompatibility groups of sweet cherry
clones. J. Amer. Soc. Hort. Sci. 92: 119—123.

Werner, D.J. and S. Chang. 1981. Stain testing viability in
stored peach pollen. HortScience 16:522-523.

15

CHAPTER II

INBREEDING, COANCESTRY, AND POUNDING CLONES OF ALHONDS

OF CALIFORNIA, MEDITERRANEAN SHORES, AND RUSSIA

l6

I W.’ev_s~~‘,,.90:: —‘ -

SUMMARY:

The cultivated almond, Prunus dulcis (Miller) D.A
Webb, has spread progressively since antiquity from along the
Mediterranean shores to the United States (U.S.), South
Africa, and Australia. To satisfy market demands and improve
cultivars, the recurrent use of four selections as parents in
the U.S. breeding programs has resulted in a mean inbreeding
coefficient (F) within the U.S. germplasm collection of 0.022.
In France, a single cultivar, 'Ferralise', has an inbreeding
value of F = 0.250, while cultivars of other almond producing
countries are noninbred (F=0). Due to the use of common
parents, the U.S., Russian, and Israeli cultivars share
coancestry, while coancestries also exist between French and
Spanish almond germplasm. Cultivars of known parentage in the
U.S., Russia, Israel, France, and Spain trace back,
respectively, to 9, 8, 3, 4, and 3 founding clones. Almond
breeding programs might, in the future, narrow the genetic

base of almond germplasm, thereby limiting genetic gain.

INTRODUCTION:

Genetic diversity of crop plants decreases the likelihood
of crop losses to insects, diseases, and unfavorable weather
conditions. Maximizing genetic diversity is also important in

breeding programs, since it maximizes the potential gain from

17

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selection. Yet, Prunus cultivars grown in the U.S. have a very
narrow genetic base. Sour cherry (g. cerasus) production in
the U.S. is a monoculture of one variety, and most commercial
peach (P. persica) varieties trace back to six parental
cultivars (Scorza et al., 1985). In almond breeding programs,
the extensive use of two cultivars, ‘Nonpareil' and ‘Mission',
suggest that the almond germplasm base may be similarly
narrow.

The cultivated almond [synonymous 2. amygdalus Batsch and
P. communis L. (Kester et al., 1991)] is believed to have
originated from the wild species Amygdalus communis (Evrinov,
1958; Grasselly and Crossa-Raynaud, 1980; Kester, 1990; Kester
et al., 1991; Kester and Asay, 1975). Amygdalus communis is
thought to be derived from hybridization among several wild
species of the subgenus Amygdalus such as E. fenzliana, P.
bucharica (Grasselly and Crossa-Raynaud, 1980; Kester and
Asay, 1975), P. ulmifolia (Evrinov, 1958; Kester and Asay,
1975), and possibly 2. kuramica (Grasselly and Crossa-Raynaud,
1980; Kester et al., 1991) and B. kotschii (C. Grasselly,
personal communication). The native habitats of the cultivated
almond are between 700 and 1700 m on the Kapet Dagh Mountains
between Iran and Russia and on the Tian Sian Mountains between
western Mongolia and Russia (Grasselly and Crossa-Raynaud,
1980; Grasselly, 1976; Kester, 1990; Kester and Asay, 1975).
Almond cultivation began during the third millennium B.C.

(Socias I Company, 1990) with sweet kerneled selections that

18

 

arose by mutation within wild populations of normally bitter
seedlings (Grasselly, 1976; Kester, 1990). Cultivated almonds
were introduced to Greece before 350 B.C. (Kester, 1990;
Socias I Company, 1990) and spread around the Mediterranean
through commercial routes (Grasselly and Crossa-Raynaud, 1980;
Kester, 1990; Kester et al., 1991; Kester and Asay, 1975;
Socias I Company, 1990). The Arabs introduced almonds into
North Africa and the Iberian peninsula during the 6th and 7th
century A.D (Kester, 1990). The introduction of almonds into
America, Australia and South Africa occurred between 1850 and
1900 (Kester, 1990). Currently almond culture is concentrated
in three main world regions: Asia, the shores of the
Mediterranean sea, and California. Limited production is found
in Australia, South Africa, Argentina, and Chile (Kester et
al., 1991).

Since antiquity, the cultivated almond has been propagated
by seed and, therefore, has been more likely to exhibit more
genetic change than a clonally propagated species would. As a
result of selection pressure, the domesticated species
progressively differentiated into separate geographical
ecotypes in the differing environments (Grasselly, 1976;
Grasselly and Crossa—Raynaud, 1980; Kester, 1990). Cultivars
were selected from seedling almond populations, grafted to
propagate desirable clones, and later established in
commercial orchards (Kester, 1990; Kester et al., 1991; Kester

and Asay; 1975, Socias I Company, 1990). Most of the leading

19

cultivars in the world originated from chance seedlings
selected from local gene pools (Kester et al., 1991).

Almond production and cultivar development followed
different patterns in different parts of the world: 1)
Seedlings and a few locally selected and vegetatively
propagated cultivars are grown commercially in Afghanistan,
Bulgaria, India, Iraq, Iran, Morocco, Pakistan, Romania,
Syria, Turkey, and Yugoslavia.

2) Active breeding and evaluation programs exist in Australia,
Greece, Israel, Italy, Spain, and Tunisia, but most of the
major cultivars originated from chance seedlings from local
ecotypes.
3) In France, the former Soviet Union, and U.S, the almond
industries rely primarily on released cultivars from breeding
programs, and old seedling or clonally selected cultivars from
landraces exist only in germplasm preservation collections.
Almond is an obligate outcrosser and susceptible to
inbreeding depression characterized by leaf abnormalities and
reduction in vigor, flower number, fruit set, seed
germination, seedling survival, and disease resistance
(Grasselly and Olivier, 1976; Grasselly et al., 1981;
Grasselly and Olivier, 1981; Socias I Company, 1990; C.
Grasselly and D. Kester, personal communication). When the
'Tuono' cultivar from the Puglia region of Italy was crossed
with unrelated cultivars, no inbreeding depression was

reported. However, when 'Tuono' was crossed with other

20

cultivars of the Puglia region or self-pollinated, inbreeding
depression occured, as expressed by low vigor and a longer
juvenile period (Socias I Company, 1990). In cultivar
development, when self-pollination or sib-mating are
practiced, the level of inbreeding in the progeny population
increases. The objective of these conservative crosses, selfs
or related crosses, is generally'to maintain the uniformity in
kernel traits required by the industry; .As a result, there is
fixation at desirable loci with an associated reduction of
fitness due to a loss of heterozygosityu The extensive use of
the 'Nonpareil' and 'Mission' cultivars in breeding programs
raises a concern of possible inbreeding depression in almond
breeding programs. The objective of the present study was to
compare the level of inbreeding, coancestry, and the genetic
contribution of founding clones among almond germplasm in

different almond producing countries.
MATERIALS AND METHODS:

Pedigrees of almond cultivars were collected from
published sources (Anonymous, 1977; Barbera et al., 1984;
Bastide and Souty, 1976; Brooks and Olmo, 1972; Brooks and
Olmo, 1982; Chessa and Pala, 1985; Costetchi, 1967; Egea et
al., 1984; Fanelli, 1939; Felipe, 1976; Felipe, 1984; Felipe
and Socias I Company, 1985; Felipe and Socias I Company, 1987;

Georgio et al., 1985; Grasselly and Crossa-Raynaud, 1980;

21

 

 

Jaouani, 1976; Kester et al., 1991; Kester et al., 1984;
Kester et al., 1985; Monastra et al., 1984; Serafimov, 1976;
Spiegel-Roy, 1976; Spiegel-Roy and Kochba, 1976a; Spiegel-Roy
and Kochba, 1976b; Spiegel—Roy et al., 1982; Stylianides,
1976; Stylianides, 1977; Vargas Garcia, 1975; Vlasic, 1976;
Wood, 1924), and breeding records. Parental relationships for
many cultivars of unknown origin have been defined by isozyme
techniques (Hauagge et al., 1987), or through pollen
incompatibility studies (Crossa-Raynaud and Grasselly, 1985;
Godini et al., 1977; Kester et al., 1985). Pedigrees for 427
almond cultivars from different almond producing countries
were obtained; however, only 124 cultivars were included in
the present study. The other 303 cultivars are of unknown
parentage. Of these 124 cultivars, 86 were American, 12
Russian, 6 Israeli, 13 French, 5 Spanish, and 3 Italian.

The inbreeding coefficient (F), given by the following
formula, is defined as the probability that 2 genes at any
locus in an individual are replicates of one and the same
gene in a previous generation. These genes are said to be

"identical by descent" (Wright, 1922).

1 nlmy
Ff}: [(3) View].

In = number of generations from one parent back to the common
ancestor.

n2 = number of generations from the other parent back to the

22

 

common ancestor.
FA = inbreeding coefficient of the common ancestor.
Estimation of the level of inbreeding by calculation of the
inbreeding coefficient gives a reasonable approximation of the
probability of fixation, even when the initial gene
frequencies are not known (Wright, 1922). Considering that
almond is an obligate outcrosser because it is self-
incompatible, all parents of unknown origin were assumed non
inbred and unrelated. The seed parent involved in an open—
pollination was also assumed to be unrelated to the pollen
parent. Inbreeding coefficients were calculated using a
computer program of Hancock and Siefker (1982).

The coancestry coefficient (CC) of perspective progeny of
2 individuals is equal to one half the covariance of the
parents. Using the same program for F calculations, the CC of
2 cultivars was calculated as F of their prospective progeny
knowing that F of an individual is equal to the CC of its
parents. The CC equals 0.500 for self-pollination, 0.250 for
parent-offspring and full-sib matings, 0.125 for half-sib
matings, and 0.063 for first cousin matings. Parentage of a
mutant of a cultivar is considered to be the same as parentage
of the mutated cultivar. Thus only the CC value of the
original cultivar is presented. However, the CC values of all
the cultivars, the mutants plus the original cultivar, were

considered for mean calculations.

23

 

The genetic contribution (GC) of a founding clone to a
cultivar was calculated as described by Sjulin and Dale

(1987):

n = number of generations in a pedigree pathway between the

founding clone and the cultivar.

x = number of pathways between the founding clone and the
cultivar.
RESULTS :

Inbreeding coefficients: Only 10 almond cultivars had

inbreeding coefficients different than zero (Table l).
Inbreeding coefficients of U.S. cultivars ranged from 0 to
0.375 (Appendix 1) with 9 of the 86 cultivars evaluated having
F>O (Table 1). The mean inbreeding coefficient for the U.S.
cultivars was 0.022. Except for the French cultivar
'Ferralise', with F=0.250, all the remaining almond cultivars
in France, Russia, Spain, and Israel are noninbred (F=0)

(Appendix 1).

Founding clones: The almond cultivars in the U.S. germplasm
collection trace back to 9 cultivars with 'Nonpareil',

24

Table 1: Country of origin, parentage, and inbreeding

coefficients for 10 almond cultivars.

 

 

Country Cultivar Presumed or Inbreeding
of reported parentagez coefficient
origin (F)
U.S. Sonora 21-19W [Nonpareil x (Nonpareil x 0.375
(5a-20) Eureka) Al—30] x 22-20
[Nonpareil x (Nonpareil x Eureka)
Al-30]
U.S. Solano 21-19W [Nonpareil x (Nonpareil x cans
(5a—3) Eureka) A1-30] x 22—20
[Nonpareil x (Nonpareil x Eureka)
Al—30]
U.S. Emerauld Mission x (Nonpareil x Mission) 0.2%
U.S. Profuse Nonpareil x Jordanolo 0.2%
U.S. Wawona Ruby x Mission 0.250
U.S. Kapareil Nonpareil x 24-6[Sel. A525 0.35
(Nonpareil x Eureka) x Eureka]
U.S. Milow Nonpareil x 24-6[Sel. A525 0.35
(Nonpareil x Eureka) x Eureka]
U.S. Vesta Solano (5a—3) x late blooming 0JB4
sport of Nonpareil
U.S. Davey Nonpareil x Sans Faute 0.063
France Ferralise Ferraduel x Ferragnes 0.2w

 

2The rule of seed parent being at left of the cross is not respected
because the direction of the cross was unknown.

25

 

'Mission', and the French 'Princesse' and 'Languedoc',
representing the highest genetic contribution (GC) (Table 2).

'Nonpareil', a seedling of 'Princesse', contributes 37.9% in

the genetic make-up of U.S. cultivars. This cultivar is
related to 65% of the cultivars studied. 'Mission' (syn.

'Texas') has a GC of 30.2% and a coancestry relationship with
49% of the cultivars under study. 'Princesse' has a GC of
21.6% , while 'Languedoc' has a GC of 14.1%. The gene pool in
California is dominated by descendants of 'Nonpareil' and
'Mission' (Hauagge et al., 1987, Kester et al., 1991).
'Nonpareil', 'Nec Plus', 'I.X.L', and 'Mission' are considered
founding clones, even though their parentage is known
(Appendix 1), because of their extensive use (mostly 'Mission'
and 'Nonpareil') in breeding programs (Appendix 2). In
addition, they originated from the first generation of
selected almonds. The maternal parent of 'Nonpareil' (as well
as of 'Nec Plus Ultra' and 'I.X.L') is believed to be a
variety known in California as 'Princesse' or 'Prince's' which
originated in the Languedoc area of France (Grasselly and
Crossa-Raynaud, 1980). The California 'Languedoc' is
apparently different from the French cultivar in the French
collection known as 'Languedoc 320' (Kester et al., 1991).
The Russian cultivars trace back to 8 founding clones
(Appendix 3, Table 2). Three of the founding clones are of
Russian origin ('Nikitski 62', 'Nikitski 1', and 'Nikitski

53'), while the 5 others are from France ('Princesse' and

26

 

Table 2: The percent genetic contribution of founding clones
over all the cultivars in their respective groups
outlined in the pedigrees in Appendices 2-6.

 

Founding clone Country of origin Genetic contribution
within each country
%

 

U.S. Germplasm:

Nonpareil U.S. 37.9
Mission U.S. 30.2
Princesse France 21.6
Languedoc France 14.1
I.X.L U.S. '3.9
Eureka U.S. 2.3
Harriott U.S. 1.6
Nec Plus U.S. 1.3
Swanson U.S. 0.7

Russian germplasm:

Nikitski 62 Russia 38.9
Princesse France 16.7
Nikitski 1 Russia 11.1
Nonpareil U.S. 11.1
Languedoc France 11.1
Nikitski 53 Russia 5.6
Fragullio Italy 5.6
Reams Italy 5.6

French germplasm:

Cristomorto Italy 41.7
Ai France 33.3
Tuono Italy 16.7
Ardechoise France 8.3

Spanish germplasm:

Tuono Italy 50.0
Ferragnes France 16.7
Tardive France 16.7

Israeli germplasm:

Greek Israel 25.0
Marcona Spain 25.0
Princesse France 25.0

 

27

'Languedoc'), Italy ('Fragullio' and 'Reams'), and U.S.
('Nonpareil'). The dominant cultivar used in Russian breeding
programs is 'Nikitski 62' (GC = 38.9%), followed by
'Princesse' (GC = 16.7%). 'Princesse', in Russia, is different
from the 'Princesse', parent of 'Nonpareil', in the U.S. and
'Poriah 10' in Israel (C. Grasselly, personal communication),
even though both cultivars are originally from France.
'Nikitski 1', 'Languedoc', and 'Nonpareil' have a CC = 11.1%.

The French almond breeding program is characterized by the
extensive use of two founding clones 'Ai' (France) and
'Cristomorto' (Italy). Both cultivars have a.(x: of 35.7%,
followed by 'Tuono' (Italy) (GC = 14.3%), and 'Ardechoise'
(France) (GC = 7.1%) (Table 2). The French cultivars released
from breeding programs trace back to 4 cultivars, 2 of Italian
origin and 2 from France (Appendix 4). All the other French
almond cultivars are of a chance seedling origin with unknown
parentage except some speculative parentage relationship
between old cultivars such as 'Fourcouronne', 'Tournefort',
and 'Tardive de la Verdiere' (Grasselly and Crossa-Raynaud,
1980).

The Spanish breeding program has released 3 cultivars to
date (Appendix 5). From this material there are 3 founding
clones: 'Tuono' from Italy (GC = 50%), 'Ferragnes' and
'Tardive de la Verdiere' from France (GC = 16.7%) (Table 2).

In Israel, 8 cultivars are of Israeli origin. Four of them

have been obtained through breeding programs (Appendix 6)

28

 

involving 3 founding clones from 3 different countries
('Greek' from Israel, 'Marcona' from Spain, and 'Princesse'
presumably from France, but probably imported from U.S.). The

GC of each of these cultivars is 25% (Table 2).

Coefficients of coancestries: Even though the inbreeding
coefficients of the almond cultivars are low (Appendix 1),
cultivars released from breeding programs show important
coancestry relationships through the repeated use of a few
superior parents.

U.S. germplasm: Coefficients of coancestries (CC) of
cultivars in the U.S. germplasm range between 0 and 0.50
(Table 3). The average CC values for individual cultivars
paired with all other cultivars range between 0 and 0.15 with
an overall mean of 0.080 for the 86 cultivars. Except for 12
cultivars of unknown origin and 'Nonpareil' and 'Mission',
which are unrelated, all of the remaining 72 cultivars (84%)
are interrelated. Seven percent of the cultivars represent a
parent-offspring relationship (CC = 0.25) while 2% are full-
sibs. Fourteen percent represent a half-sib relationship (CC
= 0.125) and 4% have CC = 0.063 (first cousin relationship).
0n the average, every cultivar is related to 38 other
cultivars with a range between 1 ('Swanson'= founding clone)
and 69 (all 'Nonpareil' x 'Mission' hybrids) (Appendix 2).

Russian germplasm: Russian cultivars have CC values between

0 and 0.250 (Table 4). The average CC value over all the

29

 

 

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31

cultivars tested is 0.061. CC mean values for cultivars
showing coancestry relationships and taken individually vary
between 0.015 and 0.103. Ten percent of the cultivars from
breeding programs have a parent-offspring relationship (CC =
0.250) and 4% are full-sibs (CC = 0.250). Fifteen percent of
these cultivars represent. a. half-sib relationship (CC =
0.125), and 5% are of first cousin type coancestry (CC =
0.063). Every released cultivar from a controlled cross is
related to at least seven other cultivars (Appendix 3).

French germplasm: The overall mean CC between cultivars
released through French breeding programs is 0.121 (Table 5).
'Ferralise' represents the highest coancestry value (CC -=
0.375) with its parents 'Ferragnes' and 'Ferraduel'. Every
cultivar is related to at least 6 other cultivars, thus
presenting a CC different than zero (Appendix 4). Eight
percent of the cultivars have a full-sib relationship and 20%
have a parent-offspring relationship (CC = 0.250). Nineteen
percent are half-sibs (CC = 0.125), and 1% are first cousins
(cc = 0.063).

Spanish germplasm: CC values for Spanish cultivars vary
between 0 and 0.250 with an overall mean of 0.108. Three
cultivars are half-sibs (CC = 0.125).The other CC's different
from zero are parent-offspring relationships (CC = 0.250)
(Table 6). Every released cultivar is related to at least 3

other cultivars (Appendix 5).

32

 

 

 

 

.ouoN I wosmmo sums poumHsomoo muumoocmoo mo mucomommmooo cmozx
.mucouoo c3ocx ozu mo weapooubcfi oc oumowoCH mozmoo»
.omnou mo umom you on muobsss um>mumoo ou Homes ombco mo oou mmouoo muonfiszu

.mmoum IH “mucouoo mm pom: mum>mumdo ammouom mo Cowmuo mo zuucsoo utomouoou mononucoumo coosuob muouuoq

 

 

 

one. .Hoocose Ha
muo. I mmoonomouo oH
mus. I I H< m
vvu. omm. I mma. monogamoum m
«we. omm. I was. omm. mccmusmo n
cos. I omm. I moo. moo. umumwuumm 6
mad. I I I mma. mma. omm. .H.ouuosoumfluo m
«we. I I omm. was. mad. mma. omm. Hmsomuumm e
and. I I omm. omm. omm. mmH. omm. omm. wmcmmuuwm m
«we. I I omm. was. was. mmH. omm. mum. mum. mmflamuumm m
mso. I I omm. moo. moo. I II was. mad. mma. mammm H
gnaw: as OH m m n o .m e m N ~H mum>flumso
.maduooum ocmoooun cm pom:
muu>mumso ocoEmo museum you momuumoocooo no mucomomuuooo "m omens

33

Table 6: Coefficient of coancestries for Spanish
almond cultivars used in breeding programs.

 

 

Cultivars 1z 2 3 4 5 6 Meanx
l A-10-6 .125 .125 .250 -Y .250 .150
2 Ayles .125 .250 - — .100
3 Moncayo .250 .250 - .150
4 Tuono(I) - - .050
5 Tardive(F) - .050
6 Ferragnes(F) .050

 

Letters 'between ‘parentheses represent country of origin of foreign
cultivars used as parents; F— France, I- Italy.

zNumbers across top of table refer to cultivar numbers at far left of
table.

yDashes indicate no inbreeding of the known parents.

xMean coefficients of coancestry calculated with dashes - zero.

 

34

Israeli germplasm: Of 11 cultivars, 9 have been involved in
breeding programs, including 3 from foreign countries
(Appendix 6). CC values vary between 0 and 0.250 (Table 7).
All CC's equalling 0.250 (31 % of the cultivars studied)
represent a parent-offspring relationship, with the exception
of a full-sib relationship between 'Solo' and 'Samish'.
Fourteen percent of the coancestry relationships are of the
half-sib type (CCI= 0.125). Every released cultivar'is related
to 5 or 6 other cultivars. Some of the original parents are
introduced cultivars. 'Princesse' is a French cultivar,
probably introduced from the U.S., 'Nonpareil' is from the
U.S., and 'Marcona' is from Spain.

Coancestrv relationships among cultivars from different
countries: Germplasm exchange between almond breeders from
different countries resulted in the common use of some
cultivars as parents. When mean CC's across cultivars within
countries are considered, U.S., Russian, and Israeli almond
cultivars share common parentage (Table 8). The mean CC
between U.S. and Russian almond cultivars is 0.030, which is
half the CC mean values within U.S. and Russian cultivars
(0.057 and 0.061, respectively). This parental relationship is
explained by the use 'Languedoc,’ old French cultivar, in
breeding programs of both countries, and from the use of
'Nonpareil' from the U.S. by Russian breeders. The mean CC
between U.S. and Israel is 0.035. The French cultivar

'Princesse' and the U.S. cultivar 'Nonpareil' are found in

35

 

Table 7: Coefficients of coancestries for Israeli almond cultivars
used in breeding programs.

 

 

Cultivars 1z 2 3 4 5 6 7 8 9 Mean"
1 Dagan .250 .125 .125 .031 .250 -Y .063 .125 .108
2 Poriah 10 - - .063 - - .125 .250 .076
3 Solo .250 .125 .250 .250 - - .111
4 Samish .125 .250 .250 - - .111
5 Kochba - .250 .250 .125 .108
6 Marcona(S) - - -_ .083
7 Greek - - .083
8 Nonpareil(U.S.) .250 .076
9 Princesse(F) .083

 

Letters between parentheses represent.country'of origin of foreign cultivars used
as parents; F- France, 8- Spain, U.S.- United States of America.

zNumbers across top of table refer to cultivar numbers at far left of table.
yDashes indicate no inbreeding of the known parents.

a‘Mean coefficients of coancestry calculated with dashes = zero.

36

 

 

Table 8: Mean of coancestry coefficients among almond
cultivars from s almond producing countries.

 

 

Country 1‘ 2 3 4 5
1 U.S.A 0.080 0.030 —Y - 0.035
2 Russia 0.061 - - 0.013
3 France 0.121 0.067 -
4 Spain 0.108 -
5 Israel 0.093

 

zNumbers across top of table refer to cultivar numbers at far left of
table.
yDashes indicate no inbreeding of the known parents.

 

37

pedigrees of cultivars from both countries. 'Princesse' and
'Nonpareil' are also the 2 common cultivars used by Russian
and Israeli breeders, resulting in a CC of 0.013 between the
2 countries. In addition, there is a coancestry relationship
among cultivars between Spain and France. This coancestry
relationship (CC‘= 0.067) resulted from the common use of the
Italian cultivar 'Tuono' in both breeding programs and by the
use of 'Ferragnes', a French release, in Spanish breeding

programs.

CONCLUSION:

The repeated use of a few founding clones and their
progeny as parents in almond breeding programs may result in
loss of genetic variability and an increase of inbreeding
depression in future generations. This is of particular
concern, since new cultivars may eventually replace the local
seedling ecotypes currently in cultivation. Similar situations
have been reported for numerous species (Hancock and Siefker,
1982; Lyrene, 1983; Martin, 1982; Mendoza and Haynes, 1974;
Reynders and Monet, 1987; Scorza et al., 1985; Sjulin and
Dale, 1987). 1

Most cultivars presently grown are F1 hybrids of
unrelated parents (e.g., Nonpareil and Mission in the U.S.).

The mean inbreeding coefficient for U.S. cultivars is lower

38

 

than that of plums (Byrne, 1989), and 4.5 to 8 times lower
than that of peaches (Reynders and Monet, 1987; Scorza et al.,
1985) . These results suggest that limited inbreeding has
occurred in U.S. almond germplasm so far. However, the high
degree of coancestry’ may limit future progress and introduce
undesirable traits (e.g, noninfectious bud-failure). The
outstanding kernel characteristics and industry importance of
'Nonpareil' led to it being used in breeding programs and
‘crossed to only a few other cultivars as donor parents for
specific traits such as late bloom from 'Mission' (Kester et
al., 1991; D. Kester personal communication). Aside from the
extensive breeding use of 'Nonpareil' and 'Mission' , these two
cultivars represent, respectively, 65% and 25% of commercial
almond production in California (Hauagge et al., 1987). This
uniformity increases the vulnerability of California almond
production to yield fluctuations due to hazards such as the
bud failure disorder that is frequent with 'Nonpareil' and its
descendants, which represent 48% of the cultivars examined
(Kester, 1969; Kester, 1970).

Almond cultivars in other countries, except the French
cultivar 'Ferralise', are noninbred. They are mostly from
chance seedlings, with a limited number of cultivars from
controlled crosses. The highest number of cultivars showing
inbreeding are from advanced breeding programs.

The major objectives in Russian breeding programs are

frost and cold resistance combined with nut quality.

39

 

'Nikitski 62', known for its late bloom.and cold hardiness, is
a frequent parent (Denisov, 1988; Rikhter, 1964; Rikhter,
1969).

In Western Europe and North Africa, the main objectives in
breeding almonds are similar, i.e. late blooming and self-
compatibility (Grasselly, 1984). As a result, only a few
common cultivars are being used extensively as parents, such
as 'Tuono', 'Ferragnes', 'Ai', and. 'Cristomorto'. Almond
breeding programs in different almond producing countries are
characterized by the use of a few superior genotypes as
parents. Clonal selection of superior genotypes, either as new
varieties or as progenitors, has provided substantial progress
in important commercial traits. The diverse almond germplasm
in several European countries is already being replaced by a
few leading and superior cultivars (Grasselly, 1984). This
situation, favored by germplasm exchange, will probably
increase coancestry relationships between released almond
cultivars of different European countries, and might, in the
future, limit genetic gain, narrow the almond genetic base,

and increase the hazard of epidemics.

40

 

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physiologiques pour l'identification de 16 variétés
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Caracteres pomologiques de 94 variétés d'amandier de la
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Bastide, J. and B. Souty. 1976. L'Amandier en France. Options
Mediterraneennes 32: 80-82.

Brooks, R.M. and H.P. Olmo. 1972. Register of new fruit and
nut varieties: Second Edition. Univ. of California Press.

pp. 1-7.

Brooks, R.M. and H.P. Olmo. 1982. Register of new fruit and
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Byrne, D.H. 1989. Inbreeding, coancestry, and founding clones
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' “56.“; _'_—‘ Rafi—2.13:1 '. ..

Fanelli, L. 1939. Varieta pugliesi di mandorle.(in Italian).
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52.

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31-38.

Felipe, A. and R. Socias I Company. 1987. 'Ayles', 'Guara',
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Georgio, V., Reina, A., and Guida, F. 1985. 'Tribuzio
Tardiva': Un semis d'amandier a floraison tres tardive.
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Godini, A., E. Ferrara, A. Reina, V. Giorgia, and F. Guida.
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G.R.E.M.P.A. CIHEAM. Bari, Italy. pp. 194-206.

Grasselly, Ch. 1976. Origine et éNolution de l'amandier
cultivé. Options Meditérranéennes 32: 45-48.‘

Grasselly, Ch. and G. Olivier. 1976. Mise en evidence de
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Grasselly, Ch., and P. Crossa-Raynaud. 1980. L'Amandier. G.P.
Maisonnueve & Larose ed.

Grasselly, Ch., P. Crossa Raynaud, G. Olivier, and H. Gall.
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Grasselly, Ch., and G. Olivier. 1981. Difficulté de survie de
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Grasselly, Ch. 1984. Réflexions diverses sur l'évolution des

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42

 

Hauagge, R., D.E. Kester, S. Arulsekar, D.E. Parfitt, and L.
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Kester, D.E. 1969. Noninfectious bud failure of almonds in
California. California Agriculture (December 1969): 12-
16.

Kester, D.E. 1970. Noninfectious bud failure from breeding
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Kester, D.E. 1990. The biological and cultural evolution of
the almond. Unpublished paper.

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Advances in Fruit Breeding. J. Janick and J.N. Moore
(eds.). Purdue Univ. Prexsster, D.E., R.N. Asay, and

W.C.Micke. 1984. 'Solano', 'Sonora', and 'Padre' almonds.
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(Prunus). Acta Hort. 290: 701-758

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variety update. Almond Board of California.

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soybean improvement program in maturity groups 00 to IV.
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37. '

43

.IIIIIIIIIIIIIIIIIIIIII-IIIIIIIIIIIIIIIZ:;;;u.“1--.-

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44

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45

Appendix 1: Parentage and inbreeding coefficients of almond
cultivars grown in the 0.8., Russia, Israel,
France, Spain, and Italy.

 

 

Cultivar Presumed or Inbreeding
reported parentagez oafifhmmnt
(F)
U.S. germplasm
Sonora (5a-20) 21-19W [Nonpareil x (Nonpareil x 0.375

Eureka) A1-30] x 22-20 [Nonpareil

x (Nonpareil x Eureka) A1-30]

21-19W [Nonpareil x (Nonpareil x 0.375
Eureka) A1-30] x 22-20 [Nonpareil

x (Nonpareil x Eureka) A1-30]

Solano (5a-3)

Emerauld Mission x (Nonpareil x Mission) 0.250
Profuse Nonpareil x Jordanolo 0.250
Wawona Ruby x Mission 0.250
Kapareil Nonpareil x 24-6[Sel. A525 0.125
(Nonpareil x Eureka) x Eureka]
Milow Nonpareil x 24-6[Sel. A525 0.125
(Nonpareil x Eureka) x Eureka]
Vesta Solano (5a-3) x late blooming 0.094
sport of Nonpareil
Davey Nonpareil x Sans faute 0.063
Arbuckle' Non pareil x Mission 0
Bigelow unknown 0
Bonita' Nonpareil x Mission 0
Ballico Mission open pollination (o.p.) 0
Belly Nec Plus mutation 0
Burbank unknown 0
Butte Nonpareil x Mission 0
Carmel' Nonpareil x Mission 0
Craven * unknown 0
Carrion Nonpareil x Mission 0
Cressey mutation of Nonpareil 0
Drake unknown 0
Dehn Northland o.p. 0
Empire Mission x peach-almond hybrid 0
Elsie' Nonpareil x Mission 0
Eureka unknown 0
Fritz' Mission o.p. 0
Golden street Languedoc o.p. 0
Godde' Nonpareil x Mission 0
Granada* Mission x IXL 0
Grace' Nonpareil x Mission 0

Parentage assuned from indirect evidence, primarily isozyme StUdieS (Hauagge et al, 1987).
Parentage from the patent description and unconfirmed.

'The rule of seed parent being at left of the cross is not respected

because the direction of the cross was unknown.

46

Appendix 1 (continued): Parentage and inbreeding coefficients of
almond cultivars grown in the 0.8., Russia

Israel, France, Spain, and Italy.

 

 

Cultivar Presumed or Inbreeding
reported parentagez coefficient

(F)
Heart' Nonpareil x Mission 0
Hoover’ Nonpareil x Mission 0
Hallshardy peach-almond hybrid 0
Harpareil Nonpareil x Harriot 0
Harriott unknown 0
Harvey' Nonpareil x Mission 0
IXL Princesse o.p. 0
Janice' Nonpareil x Mission 0
Jeffries mutation of Nonpareil 0
Jordanolo Nonpareil x Harriott 0
Jubilee' Nonpareil x Mission 0
Kern Royal mutation of Nonpareil 0
Kutsch Nonpareil o.p. 0
Livingston' Nonpareil x Mission 0
Lamarie unknown 0
Laprima Princesse o.p. 0
Leweling unknown 0
Legrand peach-almond hybrid 0
Merced* Nonpareil x Mission 0
Mission (Texas) Languedoc o.p. 0
Monterey* Nonpareil x Mission 0
Moneytree Nonpareil o.p. 0
Monarch' Mission o.p. 0
Norman' Nonpareil x Mission 0
Nonpareil Princesse o.p. 0
Nec Plus Ultra Princesse o.p. 0
Northland I.X.L o.p. 0
Pioneer peach-almond hybrid 0
Planada Tardy Nonpareil x Mission 0
Padre Mission x Swanson 0
Peerless unknown 0
Paxman' Nonpareil x Mission 0
Price Cluster' Nonpareil x Mission 0
Reinero Nonpareil o.p. 0
Ripon Tardy Nonpareil x Mission 0
Roy unknown 0
Ruby Tardy Nonpareil x Mission 0
Robson' Nonpareil x Mission 0

 

'Parentage asslmed from indirect evidence, primarily isozyme studies (Hauagge et al, 1987).

‘The rule of seed parent being at left of the cross is not respected because
the direction of the cross was unknown.

47

 

Appendix 1 (continued): Parentage and inbreeding coefficients of
almond cultivars grown in the 0.8., Russia
Israel, France, Spain, and Italy.

 

 

Cultivar Presumed or Inbreeding
reported parentagez coefficient
(F)
Sans Faute I.X.L o.p. 0
Sydney Special I.X.L o.p. 0
Smith X.L unknown 0
Standard unknown 0
Spencer Nonpareil o.p. 0
Sauret#1* Nonpareil x Mission 0
Sauret#2* Nonpareil x Mission 0
Swanson unknown 0
Tardy Nonpareil Nonpareil mutation 0
Thompson' Nonpareil x Mission 0
Tioga Tardy Nonpareil x Mission 0
Tokyo Nonpareil x Mission 0
Titan Tardy Nonpareil x Mission 0
Utah I.X.L o.p. 0
Valenta* Nonpareil x Mission 0
Walton unknown 0
Yosemite Nonpareil x Mission 0
Russian Germplasm
Bumagnoskorlupii (Nikitski 62 x Fragillio) 0
x Nonpareil
Desertnyi Nikitski 62 x Nikitski 1 0
Krimski Nikitski 53 x Languedoc 0
Miagkoskorlupii Nikitski 62 x Princesse 0
Nikitski 62 unknown 0
Nikitski 1 unknown 0
Nikitski 53 unknown 0
N.Pozdnetvetusei Nikitski 62 x Nikitski 1 0
Primorski Nikitski 62 x Princesse 0
Preanii (Nikitski 62 x Fragillio) 0
x Nonpareil
Sovietski Nikitski 62 x Languedoc 0
Yaltinski Nikitski 62 x Reams 0

'Parentage assumed from indirect evidence, primarily isozyme studies (Hauagge et al. 1987).

2The rule of seed parent being at left of the cross is not respected because
the direction of the cross was unknown.

48

 

 

Appendix 1 (continued): Parentage and inbreeding coefficients of
almond cultivars grown in the 0.8., Russia
Israel, France, Spain, and Italy.

 

Cultivar

Presumed or
reported parentagez

Inbreeding
coefficient

(F)

 

Israeli Germplasm

Dagan
Greek
Poriah 10
Solo
Samish
Kochba

French germplasm

Ai
Ardechoise
Belle d'Aurons
Ferralise
Ferragnes
Ferraduel
Ferrastar
Languedoc
Lauranne
Princesse
Steliette
Tardive de
la Verdiere

Spanish germplasm

A-10-6
Ayles
Jordan
Moncayo
Marcona

Italian germplasm

Cristomorto
Reams
Tuono

'Parentage assuned from indirect evidence, primarily isozyme studies (Hauagge et al, 1987).

Marcona x Poria 10
unknown

Princesse o.p.
Marcona x Greek
Marcona x Greek
Nonpareil x Greek

unknown

unknown

Ai o.p.

Ferraduel x Ferragnes
Ai x Cristomorto

Ai x Cristomorto
Ardechoise x Cristomorto
unknown

Ferragnes x Tuono
unknown

Ferragnes x Tuono
unknown

Ferragnes x Tuono

Tuono o.p.

unknown

Tuono x Tardive de la Verdiere
unknown

unknown
unknown
unknown

'The rule of seed parent being at left of the cross is not respected because.
the direction of the cross wasunknown.

49

000000

e
N
U!
0

 

OOOOOOOOOOOO

 

00000

000

Pedigree of 0.8. almond cultivars.
Cressey
PRINCESSE (F) Kern Royal
I Tardy Nonpareil -—-
O.P Jefferies

Appendix 2:

Ripon

U2

 

 

F

IXL

O.P
-TS

l
NEC PLUS I uta °O

NONPAREIL

 

r’
Sans

Faute

 

Utah

I
I Sydney
Northland
l mutation
O.P
Bell
Dehn

 

 

Davey

Kutsch-—l
Norman-—-
Reinero—-

Spencer-

 

Money-——-
Tree

EUREKA

 

-O.P

l

 

 

 

 

 

 

Harpareil
Jordanolo

l

Profuse

 

 

 

L

I
A5-25

I

l

I
A1-30
L

 

 

 

Lesemi:

22-20 21-19W

 

I

I .
U2 Solano Sonora

Vesta

 

 

24-6

 

I I
Kapareil Milow

 

Planada
Tioga
Titan
Ruby

monsooc (F)
9.2

Golden St.
MISSION

Q.£

 

 

 

 

I I
. Fritz Ballico
Monarch

SWANSON

 

 

Padre

 

PxA Wawona"

 

I
Empire

 

Emerauld

 

1_J
Arbuckle
Bonita
Butte
Carrion
Carmel
Elsie
Godde
Grace
Heart
Hoover
Harvey
Janice
Jubilee
Livingston
Merced
Monterey
Paxman
Price Cluster
Robson
Sauret#1
Sauret#2
Thompson
Tokyo
Valenta
Yosemite

IRA: Peachaalmond hybrid. O.P.z open pollination.
1k unnamed selection. bold: founding clones. (F): French origin.

50

 

 

Appendix 3: Pedigree of Russian almond cultivars.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

IJUWGUIHXM3-—-
(F)
Krimski
Sovietsky
'——-NIKITSKI 53
PRINCESSE(F) NIKITSKI 62-—- NIKITSKI 1
. Nikitski ,Desertnii
i i Podznetvetusei
Primorski Miagkoskorlupii
FRAGULLIO(I)
REAMS(I) l
U1. IRINPAJUIIL(II.S.)
Yaltinski L i
Preanii Bumagnoskorlupii

bold: founding clones.

(F): French origin. (I): Italian origin.

(U.S.): American origin.

51

Appendix 4: Pedigrees of French almond cultivars.

 

 

 

 

 

 

 

 

 

ARDECHOISE CRISTOMORTO (I) AI
|] [91.2
Belle d'Aurons
Ferrastar i I
Ferraduel Ferragnes TUONO(I)
Ferralise
Lauranne Stelliette

legend:
bold: founding clones.

(I): Italian origin.
O.P. :open pollination.

'52

‘\.——" ‘ —"m'-—"‘D _ 4‘

 

Appendix 5: Pedigrees of Spanish almond cultivars.

Tardive de la Tuono(I) Ferragnes(F)
1a Verdiere (F) l |

L
l o..p

Moncayo

 

 

A-10-6

ngend:
bold: founding clones.

(F): French origin, (I): Italian origin.
O.P. : open pollination.

53

Appendix 6: Pedigrees of Israeli almond cultivars.

PRINCESSE (F)

 

 

 

Nonpareil (U.S.)

 

 

 

 

 

 

O.P’
Poriah 10 MARCONA (S) GREEK
(Hanadiv) ' '
l I l
Dagan Solo Samish Kochba

Legend:
bold: founding clones.
(3): Spanish origin, (U.S): American origin.

O.P. : open pollination.

54

 

CHAPTER III

 

MORPHOLOGICAL VARIATION WITHIN COLLECTIONS OF
MOROCCAN ALMOND CLONES AND MEDITERRANEAN AND

ANERICAN CULTIVARS

55

am...

Cultivated almond Ppppps dulcis (Miller) D.A. Webb, the
second most important fruit crop in Morocco after olives, is
still propagated through seedlings by farmers to overcome
transplanting failure of grafted trees. Collections of
seedlings in southern. Morocco conducted since 1975 ‘have
resulted in the selection of 87 clones from this germplasm
planted at 3 experimental stations. Principal component
analysis (PCA) was used to quantify morphological variation
among a total of 81 selected Moroccan clones and introduced
cultivars. Moroccan selections tended to be characterized by
small leaves in comparison to foreign cultivars. Variability
for nut and kernel traits was identified. Several clones, such
as 'Ighri/13', 'Kelaa/7R', and 'B2/25Rfl have very good.nut and
kernel characteristics. However, the fruiting potential of
Moroccan selections remains limited, even though some of them
have a large number of spurs. No evidence was found of
separate ecotypes existing in the southern Moroccan almond

populations.

INTRODUCTION:

Cultivated almond was introduced to Morocco by the
Carthaginians between the 5th and 4th century B.C (El Khatib-
Boujibar, 1983) and by the Arabs during the 6th and 7th
century (Kester et al., 1991) . Almonds, the second most

important fruit tree crop in Morocco after olives, occupy

56

107,000 hectares, and comprise:73% of all Rosaceous species in
Morocco (Anonymous, 1990). Production is approximately 40,000
metric ton/year, and exportations of bitter almonds vary
between 1100 and 1300 metric ton/year (Anonymous, 1990).
Almonds are grown in Morocco in several regions under
different.environmental conditions. They are found between 500
and 2000 m elevation, where rainfall ranges from less than 100
to 800 mm, and on,a wide diversity of soils varying from deep
clay to shallow, calcareous soils. About 55% of the almond
trees grown in Morocco are seedlings, located.primarily in the
south, where this method of propagation still prevails. Five
percent of the total acreage is represented.by modern orchards
with known cultivars, mainly 'Marcona' from Spain and two
pollinizers 'Fournat de Brezenaud' from France and 'Ne Plus
Ultra' from the United States (Laghezali, 1985). Recently, 2
French cultivars, 'Ferragnes' and 'Ferraduel', have become
popular. The modern sector accounts for 80% of the total
almond production. About 40% of the almonds in Morocco have
been planted in the Rif Mountains in the north, mainly by the
Forestry Department to prevent soil erosion. 'Marcona' and
'Fournat de Brezenaud' have been the principal cultivars used
for this purpose (Laghezali, 1985). In many cases, these
cultivars have been overgrown by the rootstock, mainly
'Marcona' seedlings, due to unSuccessful grafting.
Additionally, dead 'trees Ihave been, replanted ‘with other

unknown cultivars or seedlings, making recognition of the 2

57

 

 

original cultivars difficult. This almond population
represents 5 million almond trees planted on 50,000 hectares.

The genetic variability in the Moroccan almond germplasm
is suspected to be extensive because of the broad geographic
distribution, different environmental conditions, prevalence
of seed propagation and the presence of peach-almond natural
hybrids (Barbeau and El Bouami, 1980b). For example, within
the same seedling orchard, up to a one month difference in
bloom time has been reported (Barbeau and El Bouami, 1979). In
some areas, such as Tinejdad and Goulmima in the south-east,
almond populations with a high proportion (up to 100%) of
"doubles" (nuts containing 2 kernels), are found because
double kernels have been selected by local growers. These
doubles seem. to present. no kernel deformities for some
genotypes (Barbeau.and.El Bouami, 1980a). Field collections of
almonds for late-bloom, frost resistance, and disease and
insect resistance have been carried outsince 1975 in the
south (Barbeau and El Bouami, 1979) as well as in the north
(Laghezali, 1985). A total of 87 almond clones including 11
peach-almond.hybrids were selected in the south along an east-
west axis and are growing at three different experimental
stations of the Institut National de la Recherche Agronomique
(INRA). .

The objective of the present study was to compare
introduced almond cultivars and selected Moroccan clones for

growth habit and leaf, nut and kernel characteristics using

58

Principal component analysis. Clustering of clones from
similar' collection areas 'would suggest. the existence of

different almond populations.

MATERIALS AND METHODS:

Plant material: Almond collections in the valleys of
southern Morocco since 1975 have resulted in the selection of
87 clones, collected from Errachidia (east) to Agadir (west)
(Fig. 1), which are planted at three INRA experimental
stations in the following sites: Marrakech, Errachidia, and
Meknes. Almonds at the experimental Station #1, located at
Meknes, were not irrigated. The station is under continental
climatic conditions where average annual rainfall is about 500
mm. At Station #2 at Errachidia and Station #3 at Marrakech,
almonds were irrigated; annual rainfall is less than 200 mm,
and conditions are arid.

Sixty-six clones, including one natural peach-almond
hybrid, out the 87 Moroccan selections, plus 14 introduced
cultivars and one hybrid fronm a 'Cristomorto' x 'Ardechoise'
cross were included in'the study (- Table 1). They were as
follows: 1) Meknes (Station #1), Nine Moroccan selections and
10 introduced cultivars; 2) Errachidia (Station #2), 37
lfloroccan selections including one natural peach-almond.hybrid
{and one almond hybrid from a cross between 'Cristomorto' and

'Ardechoise', and nine. introduced cultivars; 3) Marrakech

59

 

 

 

Fig. 1: Survey areas map of southern Morocco.
(from Barbeau and E1 Bouami, 1979, 1980).

EDITEBRANEA
SEA

““9“ AI Hatch-Ia

      
 

\
J

Rstu

ATLANTIC '
OCEAN
Aged:
fl
°La'you

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Ad Dumb

 

O. 300“

IMIOUIIIE

 

 

 

 

 

 

 

Logwira

6O

6..

 

 

 

Table 1: Origin and location of cultivars and clones.

 

 

 

Cultivar or clone code originz location
smathINoY
Foreign cultivars
Marcona S1 Spain 1 & 2
Desmayo 82 Spain 2
Cavaliera 11 Italy 2
Cristomorto I2 Italy 1 & 2
Tuono I3 Italy 1, 2 & 3
Ai Fl France 1
Ardechoise F2 France 1
Ferragnes F3 France 2 & 3
Fournat de Brezenaud F4 France 1 & 2
Burbank U1 U.S.A 1 & 2
Mission (Texas) U2 U.S.A 1
Thompson U3 U.S.A 2
Abiod T1 Tunisia 1
Hech Ben Smail T2 Tunisia 1
Moroccan selections

Cristo.x Ardech.hybrid M1 Morocco 2
Peach-almond hybrid.66 M2 Errachidia 2
Ksar Souk 1A Errachidia 1
Bualuzen 8A Meknes 1
Messaoud BB Meknes 1
Bl/SZ lB Errachidia 1
B1/S15 1C Errachidia 1
B1/Sl7 1D Errachidia 1
BZ/S7 1E Errachidia 1
B2/S9 1F Errachidia 1
BZ/Sll 1G Errachidia 1
B1/2L 1H Errachidia 2
B1/6BL 11 Errachidia 2
Bl/4R 1J Errachidia 2 & 3
Bl/SR 1K Errachidia 2
B1/7R 1L Errachidia 2 & 3
B1/8R 1M Errachidia 2
Bl/22R 1N Errachidia 2
BZ/8R 1P Errachidia 2
BZ/llR lQ Errachidia 2
B2/14R 1R Errachidia 2
B2/19R 1S Errachidia 2
B2/22R 1T Errachidia 2
B2/25R 1U Errachidia 2
Bl/13R 1V Errachidia 3

 

1 . . . .
Country of orIgIn for foreign cultivars and survey area for Moroccan clones.

y1: Meknes, 2: Errachidia, 3: Marrakech.

61

Table 1 (cont.): Origin and location of cultivars and clones.

 

 

 

 

Cultivar or clone code originz location
station No
Bl/lSR 1W Errachidia 3
B1/16R 1X Errachidia 3
Bl/17R lY Errachidia 3
B1/2R 12 Errachidia 3
B1/1L 1a Errachidia 3
B1/4L 1b Errachidia 3
B2/2R 1d Errachidia 3
82/19BL 1e Errachidia 3
B2/7R . 1f Errachidia 3
Hart/16 2A Gheris (Erfoud)Y 2
Hart/17 2B Gheris (Erfoud) 2
Hart/18J 2C Gheris (Erfoud) 3
Khorbat/3J 3A Ferkla (Tinejdad) 2 & 3
Khorbat/6J 3B Ferkla (Tinejdad) 3
Tizougaghine/SR 3C Ferkla (Tinejdad) 2
Kelaa/5R 4A Kelaa 2
Kelaa/7R 4B Kelaa 2 & 3
Skoura/2 ‘ 5A Skoura 2
Amekchoud/lJ SB Skoura(Amekchoud) 2 & 3
Amekchoud/3J 5C Skoura(Amekchoud) 2 & 3
Tiflit/ZR 50 Skoura(Tiflit) 2 & 3
Tiflit/ZV 5E Skoura(Tiflit) 3
Toundout/lR 5F Skoura(Toundout) 2 & 3
Toundout/BJ 5G Skoura(Toundout) 2 & 3
Toundout/8J 5H Skoura(Toundout) 2 & 3
Tiliwine/8V 5K Skoura(Tiliwine) 2 & 3
Tiliwine/8TER 5L Skoura(Tiliwine) 3
Ighri/lR 6A Taliwine 2
Ighri/13 6B Taliwine 2
Ighil Noughou 6C Taliwine 2
Ighri/12B 6D Taliwine 3
Ighri/13B 6E Taliwine 3
Ait Saoun/2V 7A Draa (Ait Saoun) 2 & 3
Ait Saoun/4V 7B Draa (Ait Saoun) 3
Ait Saoun/SV 7C Draa (Ait Saoun) 3
Ait Saoun/6V 7D Draa (Ait Saoun) 3
Ait Saoun/S3 7E Draa (Ait Saoun) 3
Agdz/lBL 7F Draa (Agdz) 2 & 3
Ircheg/2R 7G Draa (Agdz) 2 & 3
Tamkasselt/BR 7H Draa1(Tamkesselt) 2 & 3
Tinzouline/BV 7I Draa2(Tinzouline) 2 & 3
Tinzouline/SR 7J Draa2(Tinzouline) 3

 

1Country of origin for foreign cultivars and survey area for Moroccan clones.

yRegion (town).

62

(Station #3), 37 Moroccan selections and two introduced
cultivars.

gpgpgctgrs measupe : Twenty-six nut, kernel, and leaf
characters were measured in 1990 on 81 clones and cultivars at
the three stations. Nut and kernel width and tickness were
measured at the midpoint of the length, perpendicular to each
other, with the width being the larger dimension. Kernel
weight/nut weight is commonly used to describe shell hardness
(Kester and Asay, 1975). Seven additional growth habit
characters were included at Marrakech (Table 2). Five leaf and
fruit samples were collected from each selection and cultivar
for evaluation. Leaves were obtained from the middle portion
of 1-year-old shoots 25 to 30 cm long, at approximately 1.8
meters height around the tree. Four l-year and four 2-year-old
shoots were chosen at random for growth measurements following
an east-north-west-south rotation at approximately 1.8 meters
height around the tree.

Data analysis: The characters for the 81 Moroccan selections
and foreign cultivars were analyzed by principal component
analysis (PCA). In PCA, intercorrelations among variables
(components) are removed (Broschat, 1979), thus reducing the
number of ‘variables by linear: combination. of' correlated
characters into principal orthogonal axes (PC1,PC2,...,PCn)
which are not correlated (Philippeau, 1986). The first PC
represents the largest variance, followed in decreasing order

of variance values by succeeding axes PC2, PC3,..., PCn.

63

 

 

 

Table 2: Morphological characters and ratios employed in the

 

 

 

analyses.
Characters abreviations
Leaf characters:
Leaf blade length (mm) BL
Leaf blade width (mm) BW
Petiole length (mm) PetL
Vein angle (mid-vein) (dfi Vangl
Gland number anre
Serration number(over 1cm mid-limb) Snbre
Leaf width/leaf length LR

Total leaf length (mm)(leaf lenght + petiole length) II.

 

 

 

 

 

Leaf area (um?) (leaf length x leaf width) LA
Nut and kernel characters:

Nut weight (g)(in-shell) NWT
Nut length (mm) NL
Nut width (mm) NW .
Nut thickness (mm) (diameter) NTH
Kernel weight (g) KWT
Kernel length (mm) KL
Kernel width (mm) KW
Kernel thickness (mm) KTH
Kernel weight/nut weight (shell hardness) SH
Nut width/nut length NR1
Nut thickness/nut length NR2
Nut thickness/nut width NR3
Kernel width /kernel length KR1
Kernel thickness/kernel length KR2
IKernel thickness/kernel width KR3
Nut size (nm§)(nut length x nut width x nut thick.) Nvol

Kernel size (kern.length x kern.width x kern.thick) Kyol

Growth habit characters:

One-year-old shoot length (cm)
Number of laterals/l-year-old shoot
Number of nodes/l-year-old shoot

SlL
LAT
Nnodesl

Number of nodes/length unit (cm) of 1-year-old shoot Node

Two-year-old shoot length (cm)

Total number of spurs/Z-year-old shoot

S2L
Totsprs

Number of spurs/length unit (cm) of 2-year-old shoot Sprs

 

64

Therefore, the first two or three PC's usually account for a
large portion of the variance. This portion of the variance
becomes less important when there is a large number of
relatively independent variables (Daudin, 1982). PCA is used
to establish correlations between variables (characters in
this study) and to visualize the relationships of individuals
(selections and introduced cultivars in this study) in two or
three dimensional graphs. 1

PCA analyses were performed using the PRINCOMP procedure of
the SAS statistical package (SAS Institute Inc., 1985). Data

from each station were analyzed separately.

RESULTS:

At Station #1, nine Moroccan clones and 10 cultivars of
foreign origin were evaluated for 26 six leaf and nut and
kernel characters (Table 1). The first 3 principal components
accounted for 32.4%, 23.4%, and 13.7% of the total variance
respectively. Eight nut and kernel variables were highly
loaded on PC1 (Table 3). From high to low absolute values,
they were: nut size, kernel width, kernel weight, nut width,
Ikernel size, kernel length, kernel thickness/kernel width, and
nut length. All these traits had positive values except kernel
‘thickness/kernel width. PC2 included 5 nut and kernel traits,
4: of which were ratios, and a single leaf trait (Table 3).
They were: nut thickness/nut length, kernel width/kernel

length, nut width/nut length, petiole length, kernel

65

 

 

St.2 St.3

P03

St.1

 

St.3

PC2
St.2

 

St.1

St.3

P01
St.2

l component axes from PCA analysis of

 

St.1

Inpra

PC Axes
Stations

stations 1, 2, and 3 genotypes represented in figures 2, 3, and 4.

 

 

Table 3: Eigenvectors of the 3 pr

Traits
Nut

 

mnmwum ommnsumm w wunmsmnmm
O O O O I I O O O O O O O O O I O D
497.4 11 72005 03 m 045824718
2110 .Jal 1311-0 .20 . 010111100
O I I I O O I O I O O O O O 0 O
o o o o
7. II nu7.Av1. z. 151.7IKJIIIS 1. .4 .4.).U
O O O I I O O .....
o o o o
.000 u .1 1 .000230 0 221100022
0 I O I O ...... O
$000000 0&000000 0 000000000
. o o o o c o o o o o .
2114455 23630406 1 55 2 6773
0010001 0000.0110 1 IA!“ .0 .OOINIQ.
. O O I I O O 0 O O O 0 O O O
o u o o o
631 0 3906638 3 2665!»
flWZOJMZ 0111032“ 0 flMMHZOIZI
O O O O O I O O O C O O O O O O O 0
0000000 00000000 0 000000000
. o o o
1 7 983 2 1 910026 75
MAIN. 3mg1 ”.1 .121m1 1 OOOOOOMOO
O O I O O O O O I I O o O 0
oo.o.o.o.oo.oo 00000000 0 000000000
o o .
ue1nunu<3 7.7.9.:. 4.0.1. 1. nu1.o.L.9.n.2.7.1.
00.0.0.0.nw0.n.u. 00000000 0 000000000
.0 o o o o o o
5 2 0 8 26 5 2
O O O O I O O o O O O 0 O O C
oogooooooooo. ooooooom 0 000000000
. - a o
h In F
th th
Wt Wt h
d d t
he€l h .1 h m. r huh
tlw tlH 8 th tt
// 9’] S t 3 mg
5 SS S “SS 9 dll m n
S SS S 988 .m ..I 9 ee
8 [MM 9 lee lHen .Ill
.1.“ n I th D ID“ V Ila t/
hthk hkk hthk hkk :- .«eO .mahl
g tCEtCC QMtcetCC h “...-n rtaa
.1Wdu12d..l€l l1 d.1Zd.1.1 a tcelanldte
e .Ihiihh meeihiihh l lleeleior
ULUTSUTT rULUTSHTT fl “BBPVGSUTA
e h e
K s L

.0.”
-0.32
'0.22

-0.31

66

Laterals number

Length
Nodes/shoot
Nodes/cm

b. 2 year old shoot
Length
Spurs/shoot
Spurs/cm

 

PC1. PC2. and PC3 (or station 1 account (or 32.4%. 3.4%. and 13.7% of vanance between means. respecuvely.

PC1. PC2. and P0 {or smion Z acooum for 20.8%. 163%. and IS.I% of vananoe between means. respecuvely.
PC1. PC2. and PC3 for union 3 account (or am. l8.6%. and 11.8% of vanance between means. respecmely.

xflold numben are unable: highly loadm; on separaIe PC axes.

a. 1 year old shoot

Growth habit

thickness/kernel length, and kernel thickness. The separation
along PC3 was due to variation in 4 leaf characters and shell
hardness that loaded negatively, plus 2 nut variables loading
positively (Table 3). These variables, from high to low
absolute values, were: nut weight, nut thickness, leaf length,
shell hardness, leaf area, total leaf length, and leaf width.

When the cultivar and selection means were plotted on the
3 principal axes (Fig. 2), the 2 French cultivars 'Fournat'
(F4) and 'Ardechoise' (F2) had coordinates on the positive
extreme of PC1. These 2 cultivars had among the highest nut.
and kernel characteristics. On PC3, their position was
explained by their longest leaves, large leaf areas, and soft
shells (Table 4, Appendix 3).

'Marcona' ($1), a Spanish cultivar extensively planted in
modern Moroccan orchards, had a highly positive components for
all 3 axes (Fig. 2). It was distinguishable by its nut and
kernel characteristics, particularly nut.and kernel width that
were important on PC1 (Table 4). On PC2, 'Marcona' (81) was at
the positive end due to its high nut and kernel ratios,
indicating its characteristic square nut.and kernel shape, and
to its longest petioles (Table 4). 'Marcona' also had the
thickiest nut, the highest nut weight, and the hardest shells
which contributed to its maximum position on PC3 (Table 4,
Appendix 3).

Four Moroccan selections, 'Ksar Souk' (1A), 'Bl/SZ' (lB),

'Bl/Sl7' (lD), 'BZ/S9' (1F), clustered with 6 foreign

67

 

 

Fig. 2:

PC3

-0.56 '

 

“1.90 '
2.11

 

Position of PC scores of introduced cultivars and
Moroccan selections. Station #1.

Alphanumericals inside signs are abbreviations of trees and the
first digit refers to the country or area of origin (table 1).
Circles - clones. Diamonds - cultivars.

S1

 

 

 

 

 

 

 

 

 

 

 

 

 

-1.51

68

 

Table 4: Means of representative characters highly loading on PC1, PC2, and
PC3 axes for Figure 2 (Station 3 1).

 

 

 

 

Clone NHT‘ KHT 58 NL nu KL KU PETL BL 80 NR1 NR2 KR1 KRZ

Marcona 7.7 1.6 0.21 33.6 30.3 23.5 17.5 29.4 96.8 23.0 0.90 0.66 0.75 0.38
Crista. 3.4 1.1 0.35 34.8 23.4 23.3 14.6 18.0 84.0 31.4 0.67 0.46 0.63 0.29
Tuono 2.6 1.1 0.42 -32.4 24.8 22.9 14.6 24.8 94.6 27.0 0.77 0.51 0.64 0.33
A1 3.2 1.4 0.45 32.3 25.1 26.0 15.6 12.4 63.8 18.6 0.78 0.50 0.60 0.51
Ardech. 3.2 1.7 0.53 51.6 24.9 29.7 15.8 23.6 108.6 26.0 0.48 0.28 0.53 0.25
Fournat 4.9 2.0 0.40 40.4 27.3 30.5 16.6 23.6 109.2 35.6 0.66 0.43 0.55 0.48
Burbank 3.4 1.7 0.51 41.6 26.0 28.7 15.1 24.4 85.0 20.4 0.63 0.44 0.53 0.59
Mission 2.7 1.2 0.46 25.7 19.9 19.6 12.9 23.2 90.2 23.6 0.78 0.64 0.66 0.84
Abiod 3.2 1.2 0.38 31.6 23.7 23.5 14.9 09.6 79.6 22.4 0.75 0.49 0.64 0.48
Hech. 2.7 1.4 0.52 32.5 24.6 24.5 15.5 23.2 85.6 24.8 0.76 0.52 0.63 0.49
KsarSouk 5.0 1.3 0.25 33.7 24.5 23.9 13.8 23.4 79.4 22.4 0.73 0.47 0.58 0.34
Bualuzen 2.8 1.2 0.43 30.0 22.6 21.8 13.4 23.6 84.2 26.4 0.75 0.58 0.62 0.42
Messaoud 4.8 1.1 0.23 37.9 21.3 24.3 11.7 10.2 69.8 20.4 0.56 0.40 0.48 0.33
82/37 2.6 1.1 0.41 29.1 21.4 20.9 12.6 21.6 82.8 22.2 0.73 0.61 0.60 0.43
B1/SZ 6.3 1.5 0.24 38.5 24.7 28.2 14.7 14.8 59.0 20.8 0.64 0.48 0.52 0.28
81/s17 3.9 1.6 0.43 42.7 25.5 28.2 15.1 14.2 71.2 23.0 0.60 0.40 0.54 0.30
82/s9 3.4 1.4 0.41 37.9 22.9 26.6 14.0 17.4 71.2 22.8 0.61 0.43 0.53 0.32
821511 1.8 0.8 0.43 30.0 19.1 21.0 11.2 17.0 73.6 20.6 0.64 0.45 0.54 0.36
B1/S1S 3.8 1.0 0.26 34.7 20.9 23.8 12.2 21.6 83.6 20.8 0.60 0.46 0.51 0.32
Mean 3.8 1 3 0.38 35.3 23.8 24.8 14.3 19.8 82.7 23.8 0.69 0.48 0.58 0.34

 

‘Abbrevmiou are defined in Table 2.

659

cultivars, 'Hech Ben Smail' (T2) and {Abiod' (T1) from
Tunisia, 'Tuono' (13) and 'Cristomorto' (12) from Italy, 'Ai'
(F1) from France, and 'Burbank' (U1) from the U.S (Fig. 2).
The U.S. cultivar 'Mission' ('Texas') (U2) had a highly
negative component for PC1, a positive coordinate for P02, and
clusters with. 2 Moroccan selections 'Bualuzen' (8A) and
'BZ/S7' (1E) (Fig. 2). All were characterized by small nut and
kernel length, weight and width, thick nuts and kernels, and
long petioles (Table 4, Appendices 2 & 3) . 'Bualuzen' (8A) and
'BZ/S?‘ (1E) are from 2 different survey areas separated by
400 kilometers . Three Moroccan selections, 'BZ/Sll' (1G),
'Bl/SlS' (1C), and.'Messaoud' (8B), loaded.on the negative end
of PC1 and PC2 (Fig. 2). These clones represented low values
for kernel weight, thickness, and size, and high values for
kernel ratio 3 which loaded negatively on PC1 (Table 4,
Appendices 1 &2). 'BZ/Sll' (16) and 'B1/515'(1C) are from the
Errachidia region, 'while 'Messaoudl (BB) is from. Meknes
region, two regions of survey about 400 km apart (Fig. 1).

At Station #2, forty six selections and introduced cultivars
were studied (Table 1).

The first 3 PC's accounted for 58.2 95 of the total
cumulative variance with PC1, PC2, and PC3 accounting for
26.8% , 16(3%, and 15.1%, respectively. Nine nut and kernel
variables loading highly on PC1 were, in descending order of
importance, nut size, nut width, kernel size, nut weight,

kernel width, nut length, nut thickness, kernel weight, and

70

kernel length (Table 3). On PC2, 6 leaf characteristics had
high positive loadings (Table 3), and were, in descending
order, total leaf length, blade length, leaf area, leaf width,
petiole length, and gland number. On PC3, 4 different nut and
kernel ratios (Table 3) were important, in descending order:
nut width/nut length, nut.thickness/nut.length, shell hardness
(kernel weight/nut weight), and kernel width/kernel length.
The only trait that loaded negatively was shell hardness.
Kernel length loaded on absolute value slightly higher on PC3
than PC1, but its value was negative on PC3 (- 0.32) while
positive on PC1 (0.27).

'Ighri/13' (6B) is an outlier and falls separately on the
positive end of PC1 (Fig. 3). This genotype is from.Taliouine,
the area closest to Atlantic Ocean. It had the highest nut and
kernel values (Table 5). Omitting 'Ighri/13' (6B), the plot of
the selections and cultivars does not indicate any clearly
defined groups. However, except for the Italian cultivar
'Cristomorto' (I2), all the introduced cultivars at Station
#2, plus 'Cristomorto' x 'Ardechoise' hybrid (M1) and the
Moroccan natural peach-almond hybrid (M2) were located on the
;positive end of PC2, while Moroccan selections had coordinates
toward the negative end of PC2 (Fig. 3), indicating that
Moroccan selections were characterized by small leaf traits
(Table 5). The Spanish cultivar 'Marcona' (81), was on the
positive end of PC3, characterized by nut and kernel values

'lower than average, particularly' nut and kernel length,

71

 

 

 

Fig. 3: Position of PC scores of introduced cultivars and
Moroccan selections. Station #2.

Alphanumericals inside signs are abbreviations of trees and the
first digit refers to the country or area of origin (Table l).
Circles - clones. Diamonds - cultivars. Squares - hybrids.

 

 

 

 

 

 

 

 

 

 

 

 

 

259
U3
my:
r44
-d73-
d62
pcz

 

 

 

 

-LJ§1
3’80\
. 2.

 

 

 

 

 

 

 

72

Table 5: Means of representative characters highly loading on PC1, PC2, and
PC3 axes for Figure 3 (Station I 2).

 

 

 

 

 

Clone NUTz KUT SH NL NU KL KN PETL BL BU NR1 NR2 KR1

Ferragnes 5.1 1.6 0.30 37.6 28.8 27.8 18.3 21.8 97.4 35.2 0.78 0.49 0.72
Crist. X Ard. 5.1 1.6 0.31 38.1 28.1 24.9 16.8 26.6 86.6 28.8 0.73 0.48 0.67
Cavaliera 2.3 1.3 0.56 27.0 20.4 22.2 13.9 21.8 95.4 27.6 0.75 0.59 0.62
Burbank 2.5 1.2 0.48 28.7 22.4 22.2 13.7 28.0 90.8 31.4 0.77 0.59 0.61
Desmayo 3.8 1.1 0.29 33.1 21.3 23.9 12.4 24.8 101.8 24.4 0.64 0.42 0.51
Marcona 4.2 1.0 0.23 25.5 23.5 18.9 13.6 29.4 96.8 23.0 0.95 0.70 0.73
Tuono 3.2 1.2 0.38 32.2 24.9 22.3 15.1 24.8 94.6 27.0 0.77 0.52 0.67
Fournat 4.0 2.3 0.57 41.5 25.1 31.8 19.5 26.0 103.5 29.0 0.60 0.34 0.61
Cristomorto 4.7 1.4 0.30 37.2 25.7 24.3 15.5 18.3 86.3 31.0 0.69 0.46 0.63
Thompson 1.8 1.3 0.74 28.6 17.4 23.8 13.8 28.3 99.5 29.6 0.60 0.42 0.58
AxP/66 5.6 0.6 0.10 30.0 23.0 18.1 11.7 15.2 114.4 36.6 0.77 0.58 0.64
BZ/ZSR 5.6 2.9 0.53 41.6 26.7 26.6 16.3 19.0 90.5 25.0 0.64 0.45 0.61
BZ/ZZR 2.7 1.5 0.55 35.4 23.8 26.6 14.6 17.0 73.8 21.8 0.67 0.47 0.31
81/6BL 4.1 1.3 0.31 30.0 23.8 23.0 14.5 10.8 66.6 24.0 0.79 0.57 0.36
32/14R 2.9 1.8 0.63 39.9 18.6 31.3 12.2 20.4 87.4 30.6 0.47 0.40 0.32
82/19R 2.6 1.4 0.50 34.4 21.0 20.5 12.6 22.6 93.0 31.0 0.61 0.43 0.15
82/11R 2.7 1.3 0.46 32.4 20.4 22.9 12.5 14.8 69.0 20.4 0.62 0.48 0.54
B1/8R 3.0 1.3 0.45 42.0 19.2 29.1 11.7 22.0 84.4 27.8 0.45 0.39 0.40
B1122R 2.6 1.0 0.40 34.5 21.7 23.6 13.0 20.6 76.8 28.0 0.63 0.48 0.54
B1/7R 3.4 1.8 0.51 40.1 21.0 29.4 12.8 19.8 80.0 29.3 0.52 0.42 0.43
B1/5R 5.7 1.5 0.26 41.3 23.9 30.0 14.6 27.4 80.0 28.6 0.57 0.44 0.48
BZ/8R 6.2 1.7 0.27 35.8 27.8 22.6 16.9 24.0 88.0 24.2 0.77 0.53 0.74
B1/4R 8.7 1.8 0.20 48.3 27.0 32.7 14.5 29.3 82.8 31.5 0.55 0.42 0.44
81/2L 3.0 2.1 0.70 27.2 18.1 20.9 11.7 21.8 80.2 33.6 0.66 0.54 0.56
81/6BL 2.7 1.5 0.56 31.5 20.2 26.3 13.5 17.8 82.2 29.8 0.64 0.47 0.51
Hart/16 4.1 2.4 0.59 37.3 25.0 26.4 14.6 21.5 101.0 31.8 0.67 0.43 0.55
Hart/17 5.1 1.5 0.29 36.4 22.9 26.9 13.8 15.4 62.6 22.6 0.62 0.50 0.51
Khorbat/3J 4.2 1.1 0.26 32.8 21.3 22.8 13.6 13.0 67.4 18.8 0.65 0.47 0.59
Tizoug./5R 4.5 1.3 0.29 33.6 23.3 22.3 14.6 20.8 74.4 25.8 0.69 0.54 0.65
Kelaa/5R 3.5 0.9 0.25 30.3 21.0 20.7 13.1 24.4 95.4 25.2 0.69 0.50 0.63
Kelaa/7R 6.5 1.5 0.24 33.9 27.3 22.7 15.9 27.0 98.0 30.0 0.80 0.61 0.69
Skoura/2 2.3 1.4 0.59 35.5 21.1 24.6 13.8 17.5 88.3 28.9 0.59 0.41 0.56
Tiliwine/8V 3.2 1.6 0.49 34.8 21.0 26.2 14.2 16.0 83.4 25.8 0.60 0.43 0.54
Tiflit/ZR 6.0 1.5 0.25 41.3 26.8 27.3 15.6 20.8 74.8 24.2 0.65 0.43 0.57
Toundout/3J 2.8 0.8 0.26 29.8 20.3 21.8 13.1 26.0 79.0 26.6 0.68 0.50 0.60
Toundout/1R 6.6 2.0 0.29 39.2 25.5 26.7 15.9 18.6 80.8 25.8 0.65 0.45 0.59
Toundout/8J 2.0 1.3 0.62 32.5 16.2 24.7 12.4 20.0 98.2 26.2 0.49 0.44 0.50
Amekchoud/1J 4.6 1.5 0.30 33.9 23.1 23.5 14.2 20.0 83.4 28.6 0.68 0.58 0.60
Amekchoud/3J 7.4 1.9 0.26 40.1 27.5 28.6 16.8 15.5 82.0 26.3 0.69 0.50 0.59
Ighri/13 9.9 3.0 0.30 47.8 33.7 32.8 19.1 24.0 76.3 20.3 0.70 0.49 0.57
Ighri/1R 4.8 '1.5 0.31 32.7 25.4 22.8 13.6 20.7 86.3 27.3 0.77 0.61 0.59
Ighil Noughou 5.1 1.4 0.27 32.8 23.9 23.8 14.6 16.0 65.2 19 6 0.72 0.55 0.61
Ait Saoun/ZV 4.8 1.4 0.29 37.3 24.2 26.4 13.2 17.4 83.2 29.2 0.64 0.49 0.50
Agdz/1BL 3.9 1.3 0.34 30.0 22.5 25.4 14.0 23.2 81.6 29.8 0.77 0.54 0.32
Ircheg/ZR 2.8 0.7 0.25 26.7 20.3 18.7 11.9 13.8 68.2 21.8 0.75 0.58 0.63
Tinzouline/3V 2.8 1.2 0.41 26.6 20.9 20.8 14.0 16.2 63.2 20.4 0.78 0.58 0.45
Mean 4.2 1 5 0.39 34.8 23.2 24.8 14.3 20.9 84.7 27.0 0.68 0.50 0.60

 

zAbbn-vialiom are defined in Table 2.

73

inducing high nut and kernel ratios (Table 5). On the negative
side of PC3 were found two Moroccan selections, 'B1/8R' (1M)
and '82/14R' (1R), because of their long and small size nuts
and kernels (Table 5).

At Station # 3, 37 Moroccan clones and 2 foreign cultivars,
'Tuono' (I3) from Italy and 'Ferragnes! (F3) from France, were
studied (Table 1). PC1, PC2, and PC3 represented 56% of total
variance with respectively 25.4% , 18.6% , and 11.8% (Table
3). PC1 included variation for'7rnnzand kernel traits, which,
in descending order of importance, were nut size, nut width,
nut weight, nut thickness, kernel width, kernel size, and
kernel weight. On PC2 the highly loaded variables were 6 nut
and kernel traits (Table 3), mainly nut and kernel ratios, and
included, from high to low absolute value: nut thickness/
length, kernel thickness/length, kernel length, nut length,
nut width/length, and kernel width/kernel length . Nut length
and nut width were loaded negatively. Six growth habit traits
loaded on PC3 (Table 3). They were from high to low values:
number of laterals/l-year-old shoot, number of nodes/l-year-
old shoot, number of spurs/cm of 2-year-old shoot, number of
spurs/Z-year-old shoot, 1-year-old shoot length, and 2-year-
old shoot length. The 2 spur variables were loaded positively
while vegetative growth variables were loaded negatively,
suggesting that l-year-old shoot growth and growth on 2-year-
old shoots were inversely related (Table 6). The leaf traits

were of minor importance, only loading on PCS, which

74

represented 7.9% of the total variance.

Independent clustering groups were not obtained (Fig. 4) .
'Kelaa/7R' (4B), the only clone from the Dades region (Fig. 1)
was at the extreme positive side of PC1 and PC2 (Fig. 4). It
had values above average for all nut and kernel characters
(Table 6), including the highest values for nut width and
kernel width. On PC2 it was characterized by the highest nut
width/length, nut thickness/length, and kernel width/length
(2nd value after 'Tinzouline/BV' (71)).

Nineteen Moroccan selections, 'Tinzouline/BV' (7I),
'Tinzouline/SR' (7J), 'Tamkasselt/BR' (7H), 'Ircheg/ZR' (7G),
'Ait Saoun/ZV' (7A), 'Ait Saoun/6V' (7D), 'Ait SaOun/S3' (7E),
'Agdz/IBL' (7F), 'Amekchoud/lJ'(SB), 'Tiflit/ZR' (SD),
'Tiflit/ZV” (5E), 'Toundout/lR' (5F), “Toundout/BJ' (5G),
'Bl/7R' (1L), 'Bl/lSR' (1W), 'Bl/l6R' (1X), 'BZ/ZR' (1d),
'Khorbat/3J' (3A) , and 'Hart/18J' (2C) , loaded on the negative
side of PC1 and the positive side of P02 (Fig. 4), and were
characterized by small nuts and kernels, high nut and kernel
ratios, and a high number of spurs/cm of shoot length, even
though shoot growth was variable among selections (Table 6).
The first eight selections are from the Draa valley survey
region [Ait Saoun, Agdz, Tamkasselt, Tinzouline. (Fig. 1)].
They represent 8 of 10 genotypes tested from this region.

'Bl/4R' (1J), 'Bl/ZR' (12), 'Bl/lL' (1a) 'Bl/4L' (1b),
'82/19BIH (1e), and 'BZ/7R' (1f), were on the negative side of

PC2 (Fig. 4) because of their low nut and kernel ratios, due

75

 

Fig. 4:

PC3

1.76

-0.03

“1.81 ‘

 

-....o J
3.14

 

Position of PC scores of introduced cultivars and
Moroccan selections. Station #3. -

Alphanumericals inside signs are abbreviations of trees and the
first digit refers to the country or area of origin (Table l).
Circles - clones. Diamonds - cultivars.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

-l.63

76

 

Table 6: Means of representative characters highly loading on PC1, PC2, and

PC3 axes for Figure 4 (Station I 3).

 

KR1 KRZ

SPRS NR1 NR2

1101‘ m1 m. 1111 KL Kw LAT s1L SZL

cum

 

 

62781118M488462894164062028543316426028
3322333J.2 5523333453433532553543333443
O O O O I O I O O ..... O O O O C O O o O I I I O O O .
000000000000000000000000000000000000000
705977890072135547298283 4663820086585
654454545546546556664645w5556456555656fl
O O I O O O O O I O I O I I ...... O O O O O O I O O O O O O 0 O O I O
000000000000000000000000000000000000000
252M4“736154056590650294421972317334446
I O O O I O O O 0 O O o O I 0 I I O I O D O O O O O O O O O D O O D
000000000000000000000000000000000000000
1.5111 9 1788 55 7 82 7906 637 46 S
”65666nw5 5554nw55nfl5 55 £5675&766M77fl8
o a a o a a a a a a a o a a a a a a o
000000000000000000000000000000000000000
93093370109078673812785138856 500360441

430.15320.530.320330221Ifl3.1342142 0.146423le

0.0.0.0.0.0.0.0.00.00000000000000000000.000000000

0839519‘74270704872679688012919109026853

5019401293104157964200738112098238093162

1121.11.11 111111.111 31122111.. 1111! 1.1111

2682AJ3857837028216221362626281232409011

LahnU.1ath.~I.62~/38390018297790105638178576678

2000000150880000020640020002060000004000

000000000000000012010000000000000000000

1525427756964845954558463.002880437524896

540222119312313106240002335451341123225

11111111 111111111111111111111111111111

3951869605131.16515195335295003805380972

2825259 9650670004011713246554442910191

Lag/5222131222222222222122222222222122212

al al.-5
4381019262991907889596671
121121211112

22.0

3248248791233
2212221212112 2221221112222
282563583126329640472085399165881398727
I O O O O O I ........... O O O O ..... O ...... O
273425737478787983769370140242419698906
333333232337.332223232223334333332222232

26517764321035988117491776481058158.0568
342323221422332225352112325452342223332

RRV
JJ VVVV3 353
13 JJRV 24565 [I]

s 36 VR/IIIEBB mmm Rlln
JI/R ZZttt T23 Lze.‘..l
L art? Ill. mm 811 COOS/Slit
QBRR n 1.“.MIhhtt .‘llI/aaaaaelgsww

a935R RRmeL/ acc.1l..l "H.35ISSSSS/ea

r111411221 [wtrrakkll ..I..Irr. Z." 22
ez11111211121ahhe ..l.‘ ..I.‘lgg.ll.al.el.ul.1lgr ..l.‘
TFBBBBBBBBBBBBRKKK TclvlvlTvI-IIIAAAAAAITTI

0.30 0.67 0.47 0.56 0.34

13.5

22.8 12.8 0.2 14.3

31.7 21.0

15

32

Mean

 

zAl'zlarcvialiom are defined In Table 2.

77

to their long nuts and kernels (Table 6) . All these selections
are from the Errachidia area of survey (Fig. 1).

The 'Ferragnes' (F3) cultivar from France and 'Tuono'
(13) cultivar from Italy, plus 5 Moroccan selections
'Ighri/lZB' (6D), 'Ighri/lBB' (6E), 'Amekchoud/BJ' (5C),
'Tiliouine/8TER' (5L), and 'Ait Saoun/SV' (7C), loaded
positively on PC1 and PC3 (Fig. 4), and were characterized by
high nut and kernel weights, with a tendency to large nuts and
kernels (Table 6). 'Tuono' and 'Ferragnes' had vigorous shoot
growth and high spur production, while Moroccan selections
with vigorous shoot growth had low spur production [i.e.,
'Amekchoud/3J' (5C) and 'Ait Saoun/SV'(7C)] (Table 6). Other
remaining selections of the group, excluding 'Ighri/lZB' (6D)
had average shoot growth and a tendency to low spur production
(Table 6).

'Khorbat/6J' (38) had the most negative value on PC3 (Fig.
4) because of its highest values for the number of laterals
and for 1 and 2-year-old shoot growth, and its low spur number
(Table 6).

About one third of the selections at Station 3 had better
or equivalent spur production potential relative to foreign
cultivars. Among these, 'Ait Saoun/ZV' (7A), 'Ait Saoun/SB'
(7E), 'Bl/lSR' (1W), 'BZ/ZR' (ld), 'Tinzouline/BV' (71),
'Tiflit/ZV' (5E), and 'Ighri/lzB' (60) showed high spur
production. However, all of these selections, except

'Ighri/12B' (6D), had small nuts and kernels.

78

 

 

CONCLUSION:

Nut and kernel, leaf, and growth habit traits that were
consistently highly loaded on the three principal axes at the
three stations were nut and kernel weight, nut and kernel
sizes, explained, respectively, by high loading of nut length
and. width, and, kernel length. and. width, nut. and. kernel
‘width/length. and. thickness/length. s, Iblade length, blade
width, petiole length, total leaf length, leaf area, one and
two-year-old shoot length, number of laterals and number of
nodes/cm on 1-year-old .shoots, number of spurs/é-year-old
shoot, and number of spurs/cm on 2-year-old shoots. Nut and
kernel measurements were always highly loading on PC1, and nut
and kernel ratios on PC2, except for Station 2 where they
loaded on PC3. When growth habit was not considered, leaf
traits loaded either on PC2 or PC3. Growth habit traits loaded
on PC3 at Station 3. For all 3 analyses, nut and kernel traits
always explained the largest portion of the variance,_
indicating that these traits were more variable than growth
habit and leaf traits. Growth habit traits, measured at
Station 3 only, were more important than leaf traits in
explaining variation among the selections.

- Station 1, which had the largest number of foreign
cultivars (10 cultivars), with cultivars to clone ratio of
approximately 1:1, presented greater differences among trees
in the morphological traits studied than the other stations.

From this morphological study, there was no evidence that

79

 

Moroccan almond ecotypes exist in this collection, since
clones did not cluster by survey area. The absence of separate
ecotypes could be explained by the fact that all surveyed
areas, as shown in Fig. 1, are on the main southern road
connecting the east and the west of the country and that seeds
were spread by continuous population flow within the survey
axis. However, at Station 3, most of the selections from the
Draa Valley were characterized by small nuts and kernels and
high spur production and clustered together. The Draa valley
separates from the main east-west axis at Ouarzazate toward
the south, and population flow is less important there. At
Station 3 also, most of the selections from Errachidia survey
area, south east of Morocco, clustered together, and were
characterized by long nuts and kernels.

Some foreign cultivars, even 'though clustering"with
Moroccan clones and other cultivars, remained grouped with
respect to their country of origin. Most foreign cultivars
were distinguished from the majority of the Moroccan
selections by larger leaves and softer shells (kernel
weight/nut weight). The tendency of Moroccan clones to have
small leaves and petioles could have resulted from natural
selection for resistance to drought conditions. In‘ most
surveyed areas, almond trees are rarely irrigated. Hard shells
have been selected by the Moroccan growers to prevent insect
damage during storage.‘ Almond is preferred to other fruit

crops because it is not a perishable commodity and there are

80

 

no special conditions for storage when it has hard shells.
Except for tendencies toward small leaves and hard shells, nut
and kernel characteristics were highly variable ranging from
nut and kernels with characteristics better or similar to good
foreign cultivars to small nut and kernels with little

commercial value.

In summary, although the morphological variation observed
among Moroccan selections did not suggest the existence of
separate ecotypes, morphological differences were identified
between Moroccan selections and introduced cultivars. Eleven
clones were identified that are presumably adapted to dry
conditions with good nut and kernel quality ['Ighri/13' (6B),
'Ighri/lZB' (6D), 'B2/25R' (1U), and 'Kelaa/7R' (48)] and/or
have good yield potential ['Ait Saoun/ZV' (7A), 'Ait Saoun/SB'
(7E), 'Bl/lSR' (1W), 'BZ/ZR' (1d), 'Tinzouline/3V' (71),
'Tiflit/ZV' (SE), and 'Ighri/lZB' (6D)). These selections
could be used as parents in a breeding program, or directly as

new cultivars.

81

 

Literature Cited

Anonymous. 1990. Secteur amandier. Bilan de la campagne 1989-
1990. Division de l'Horticulture. Direction de la
Production Végétale. Ministere de l'Agriculture et de la
Réforme Agraire. pp. 1-3.

Barbeau ,G., and A. El Bouami. 1979. Prospection de tardivité
de floraison chez l'Amandier dans le sud Marocain. Fruits
34 (2): 131-137.

Barbeau ,G., and A. El Bouami. 1980a. Prospections "Amandier"
dans le sud Marocain. Fruits 35 (1): 39-50.

Barbeau ,G., and A. El Bouami. 1980b. Les hybrides amandier x
pecher naturels du sud Marocain. Fruits 35 (3): 171-176.

Broschat ,T.K. 1979. Principal component analysis in
horticultural research. HortScience 14: 114-117.

 

Daudin, J. 1982. Analyse en composantes principales.
Mathematique et Informatique Document. Institut National
Agronomique. Paris Grignon. pp. 1-63.

EL Khatib-Bouj ibar, N. 1983. Le Maroc et Carthage. Le Memorial
du Maroc (I). Nord Organisation ed. p. 140.

Kester, D. E., T. M. Gradziel, and Ch. Grasselly. 1991.
Almonds (Prunus). Genetics Resources of Temperate Fruits
and Nut Crops. Acta Hort. 290: 701—758.

Laghezali, M. 1985. L'Amandier au Maroc. Options
Mediterraneennes 85 (1): 91-96.

Phillipeau, G. 1986. Comment interpreter les résultats d'une
analyse en composantes principales. ITCF document.
Institut.Technique des Céréales.etmdes Fourrages. pp. 1-28.

SAS Institute, Inc. 1985. SAS user's guide: Statistics. 5th
edition. SAS Institute, Inc., Cary, N.C.

82

Appendix 1: Means of characters measured highly loading

on PC1 axis for Figure 2 (station # 1).

 

 

Clone NL NW KL KW KWT NVOL KVOL KR3z
Marcona 33.6 30.3 23.5 17.5 1.6 22454 3652 0.51
Cristo. 34.8 23.4 23.3 14.6 1.1 13040 2348 0.47
Tuono 32.4 24.8 22.9 14.6 1.1 13276 2537 0.52
AI 32.3 25.1 26.0 15.6 1.4 13054 3212 0.51
Ardech. 51.6 24.9 29.7 15.8 1.7 18414 3395 0.46
Fournat 40.4 27.3 30.5 16.6 2.0 18999 4055 0.48
Burbank 41.6 26.0 28.7 15.1 1.7 19850 3870 0.59
Mission 25.7 19.9 19.6 12.9 1.2 8627 2825 0.84
Abiod 31.6 23.7 23.5 14.9 1.2 11602 2538 0.48
Hech. 32.5 24.6 24.5 15.5 1.4 13425 2889 0.49
KsarSouk 33.7 24.5 23.9 13.8 1.3 13019 2662 0.59
Bualuzen 30.0 22.6 21.8 13.4 1.2 11938 2680 0.68
Messaoud 37.9 21.3 24.3 11.7 1.1 12078 2277 0.68
B2/S7 29.1 21.4 20.9 12.6 1.1 11133 2429 0.72
Bl/SZ 38.5 24.7 28.2 14.7 1.5 17607 3212 0.53
Bl/Sl7 42.7 25.5 28.2 15.1 1.6 18471 3642 0.56
BZ/S9 37.9 22.9 26.6 14.0 1.4 14228 3148 0.61
BZ/Sll 30.0 19.1 21.0 11.2 0.8 7828 1807 0.68
Bl/SlS 34.7 20.9 23.8 12.2 1.0 11534 2201 0.63
Mean 35.3 23.8 24.8 14.31 1.3 14242 2917 0.58

 

zVariable loading negatively on PC1.

83

 

Appendix 2: Means of characters measured highly loading

on P02 axis for Figure 2 (station # 1).

 

 

Clone FRl FRZ KR1 KR2 KTH PETL
Marcona 0.90 0.66 0.75 0.38 8.92 29.4
Cristomorto 0.67 0.46 0.63 0.29 6.82 18.0
Tuono 0.77 0.51 0.64 0.33 7.58 24.8
Ai 0.78 0.50 0.60 0.51 7.94 12.4
Ardechoise 0.48 0.28 0.53 0.25 7.26 23.6
Fournat 0.66 0.43 0.55 0.48 7.98 23.6
Burbank 0.63 0.44 0.53 0.59 8.92 24.4
Mission 0.78 0.64 0.66 0.84 10.84 23.2
Abiod 0.75 0.49 0.64 0.48 7.22 9.6
Hech B.Smail- 0.76 0.52 0.63 0.49 7.60 23.2
Ksar Souk 0.73 0.47 0.58 0.34 8.06' 23.4
Bualuzen 0.75 0.58 0.62 0.42 9.12 23.6
Messaoud 0.56 0.40 0.48 0.33 7.96 10.2
BZ/S7 0.73 0.61 0.60 0.43 9.08 21.6
Bl/SZ 0.64 0.48 0.52 0.28 7.76 14.8
Bl/Sl7 0.60 0.40 0.54 0.30 8.44 14.2
BZ/S9 0.61 0.43 0.53 0.32 8.56 17.4
BZ/Sll 0.64 0.45 0.54 0.36' 7.64 17.0
Bl/SlS 0.60 0.46 0.51 0.32 7.58 21.6
Mean 0.69 0.48 0.58 0.34 8.17 19.8

 

84

 

Appendix 3: Means of characters measured highly loading

on PC3 axis for Figure 2 (Station # 1).

 

 

Clone BLz BWz LLz LAz NWT NTH SH

Marcona 96.8 23.0 126.2 2227 7.72 22.06 0.21
Cristo. 84.0 31.4 102.0 2631 3.38 15.92 0.35
Tuono 94.6 27.0 119.4 2554 2.64 16.48 0.42
AI 63.8 18.6 76.2 1195 3.20 16.10 0.45
Ardech. 108.6 26.0 132.2 2824 3.22 14.24 0.53
Fournat 109.2 35.6 132.8 3927 4.88 17.48 0.40
Burbank 85.0 20.4 109.4 1734 3.44 18.22 0.51
Mission 90.2 23.6 113.4 2139 2.70 16.54 0.46
Abiod 79.6 22.4 89.2 1780 3.20 15.46 0.38
Hech. 85.6 24.8 108.8 2137 2.68 16.78 0.52
Ksar Souk 79.4 22.4 102.8 1786 5.02 15.76 0.25
Bualuzen 84.2 26.4 107.8 2241 2.82 17.54 0.43
Messaoud 69.8 20.4 80.0 1421 4.82 14.96 0.23
BZ/S7 82.8 22.2 104.4 1839 2.64 17.80 0.41
Bl/S2 59.0 20.8 73.8 1230 6.32 18.40 0.24
Bl/Sl7 71.2 23.0 85.4 1639 3.86 16.72 0.43
BZ/S9 71.2 22.8 7 88.6 1631 3.40 16.40 0.41
BZ/Sll 73.6 20.6 90.6 1517 1.78 13.62 0.43
81/815 83.6 20.8 105.2 1740 3.80 15.90 0.26
Mean 82.7 23.8 102.5 2010 3.76 16.65 0.38

 

zVariables loading negatively on PC3.

85

 

Appendix 4: Means of characters censured hidaly loading
on P61 axis for Figure 3 (Station I 2).

 

 

 

 

Clone NUT ML NU NTH K01 KL KN NVOL KVOL
Ferragnes 5.12 37.62 28.82 17.36 1.62 27.78 18.28 19588 4502
Grist. X Ard. 5.14 38.08 28.08 18.44 1.62 24.90 16.80 19695 4086
Cavaliera 2.28 27.04 20.36 16.18 1.28 22.24 13.88 8914 3072
Burbank 2.48 28.72 22.36 17.00 1.18 22.24 13.66 10949 2818
Desmayo 3.80 33.10 21.28 14.04 1.08 23.90 12.40 10004 2325
Marcona 4.18 25.54 23.48 17.30 0.96 18.86 13.64 10392 2351
Tuono 3.24 32.16 24.90 16.94 1.22 22.32 15.10 13592 2732
Fournat 3.95 41.45 25.05 14.30 2.25 31.80 19.45 14685 5723
Cristomorto 4.68 37.20 25.70 17.23 1.43 24.28 15.50 16513 3277
Thompson 1.80 28.58 17.38 12.03 1.33 23.75 13.83 6081 2743
AxP/66 5.56 29.58 23.00 17.18 0.58 18.12 11.72 11729 1454
BZ/ZSR 5.63 41.63 26.73 19.05 2.88 26.60 16.28 21585 3539
32/22R 2.70 35.36 23.84 16.68 1.50 26.62 14.58 14081 3223
B1/6BL 4.08 30.04 23.80 17.20 1.26 22.98 14.54 12329 2837
BZ/14R 2.90 39.84 18.56 16.02 1.84 31.32 12.18 11869 3918
82/19R 2.64 34.44 21.04 14.98 1.36 20.47 12.58 10898 2145
82/11R 2.68 32.42 20.38 15.80 1.26 22.86 12.50 10601 3014
B1/8R 2.96 41.96 19.18 16.38 1.34 29.10 11.74 13199 2979
B1/22R 2.56 34.46 21.68 16.64 1.02 23.62 12.98 12464 2233
B1/7R 3.43 40.08 21.00 17.05 1.78 29.40 12.75 14443 3697
B1/5R 5.68 41.30 23.88 18.50 1.48 30.02 14.60 18266 3238
BZ/8R 6.22 35.78 27.84 19.22 1.72 22.62 16.90 19279 4064
B1/4R 8.65 48.33 26.98 20.73 1.75 32.68 14.53 27165 3717
B1/ZL 3.04 27.24 18.08 14.84 2.14 20.90 11.72 7307 1838
81/68L 2.70 31.54 20.16 14.84 1.54 26.26 13.48 9533 3416
"art/16 4.13 37.25 25.00 16.05 2.40 26.38 14.60 14942 2852
Hart/17 5.14 36.36 22.88 18.22 1.54 26.88 13.84 15188 3142
Khorbat/3J 4.16 32.80 21.34 15.62 1.12 22.76 13.58 11065 2374
Tizoug./5R 4.52 33.56 23.34 18.14 1.34 22.30 14.58 14318 3167
Kelaa/5R 3.50 30.32 20.98 15.22 0.90 20.74 13.14 9818 1981
Kelaa/7R 6.45 33.90 27.25 20.68 1.53 22.70 15.85 19220 3608
Skoura/2 2.25 35.53 21.08 14.83 1.35 24.63 13.83 11260 3117
Tiliwine/8V 3.24 34.82 20.98 15.20 1.58 26.18 14.22 11266 3208
Tiflit/ZR 6.02 41.24 26.84 17.78 1.54 27.34 15.64 19757 3121
Toundout/3J 2.82 29.82 20.32 14.88 0.80 21.80 13.12 9106 2390
Toundout/1R 6.60 39.22 25.54 17.88 1.96 26.68 15.92 17935 3861
Toundout/8J 2.04 32.46 16.22 14.54 1.26 24.72 12.44 7641 ‘ 2419
Amekchoud/1J 4.60 33.88 23.14 19.64 1.46 23.50 14.18 15419 3195
Amekchoud/3J 7.35 40.12 27.53 19.45 1.85 28.60 16.75 21136 4428
Ighri/13 9.90 47.80 33.73 23.50 3.00 32.83 19.07 38006 7448
19hri/1R 4.77 32.70 25.43 19.90 1.50 22.80 13.57 16531 3524
Ighil Noughou 5.12 32.84 23.90 18.20 1.42 23.80 14.56 14417 3400
Ait Saoun/ZV 4.80 37.32 24.22 18.54 1.42 26.36 13.24 16864 3301
Agdz/1BL 3.86 30.00 22.46 _ 15.74 1.34 25.36 14.04 10662 2964
Ircheg/ZR 2.76 26.68 20.26 15.46 0.70 18.66 11.88 8449 1493
Tinzouline/3V 2.82 26.62 20.90 15.60 1 16 20.80 14.04 8672 2776
”can 4.23 34.75 23.19 16.98 1.49 24.81 14.30 14279 3189
86

Appendix 5: Means of character: named hidily loading
on PC2 axis for Figure 3 (Station I 2).

 

 

 

Clone BL BU PETL LL GNBRE LA

Ferragnes 97.40 35.20 21.80 119.20 4.20 3427
Crist.x Ard. 86.60 28.80 26.60 113.20 3.60 2500
Cavaliera 95.40 27.60 21.80 117.20 3.20 2632
Burbank 90.80 31.40 28.00 118.80 6.80 2856
Desmayo 101.80 24.40 24.80 126.60 4.40 2486
Marcona 96.80 23.00 29.40 126.20 3.80 2227
Tuono 94.60 27.00 24.80 119.40 5.20 2554
Fournat 103.50 29.00 26.00 129.50 5.50 3006
Cristomorto 86.25 31.00 18.25 104.50 3.00 2671
Thompson 99.50 29.25 28.25 127.75 5.25 2910
AxP/66 114.40 36.60 15.20 129.60 1.80 4210
BZ/ZSR 90.50 25.00 19.00 109.50 1.75 2267
BZ/ZZR 73.80 21.80 17.00 90.80 3.40 1624
B1/6BL 66.60 24.00 10.80 77.40 2.00 1601
82/14R 87.40 30.60 20.40 107.80 3.80 2722
82/19R 93.00 31.00 22.60 115.60 3.20 2857
82/11R 69.00 20.40 14.80 83.80 1.80 1415
81/8R 84.40 27.80 22.00 106.40 2.40 2363
B1/22R 76.80 28.00 20.60 97.40 3.20 2150
81/7R 80.00 29.25 19.75 99.75 3.25 2337
B1/5R '80.00 28.60 27.40 107.40 2.60 2285
32/8R 88.00 24.20 24.00 112.00 4.20 2123
B1/4R 82.75 31.50 29.25 112.00 3.50 2609
81/2L 80.20 33.60 21.80 102.00 3.20 2715
81/6BL 82.20 29.80 17.80 100.00 3.20 2450
Hart/16 101.00 31.75 21.50 122.50 3.25 3210
Hart/17 62.60 22.60 15.40 78.00 3.20 1432
Khorbat/3J 67.40 18.80 13.00 80.40 1.40 1268
Tizoug./5R . 74.40 25.80 20.80 95.20 2.00 1921
Kelaa/5R 95.40 25.20 24.40 119.80 4.80 2419
Kelaa/7R 98.00 30.00 27.00 125.00 5.50 2954
Skoura/2 88.25 28.75 17.25 105.50 3.00 2548
Tiliwine/8V 83.40 25.80 16.00 99.40 3.00 2166
Tiflit/ZR 74.80 24.20 20.80 95.60 2.80 1811
Toundout/3J 79.00 26.60 26.00 105.00 3.40 2102
Toundout/1R 80.80 25.80 18.60 99.40 3.40 2084
Toundout/8J 98.20 26.20 20.00 118.20 4.00 2585
Amekchoud/1J 83.40 28.60 20.00 103.40 5.00 2390
Amekchoud/3J 82.00 26.25 15.50 97.50 2.25 2164
Ighri/13 76.33 20.33 24.00 100.33 7.00 1552
lghri/1R 86.33 27.33 20.67 107.00 5.00 2388
Ighil Noughou 65.20 19.60 16.00 81.20 3.20 1276
Ait Saoun/ZV 83.20 29.20 17.40 100.60 4.00 2434
A9d2/18L 81.60 29.80 23.20 104.80 3.40 2445
lrcheg/ZR 68.20 21.80 13.80 82.00 2.00 1478
Tinzouline/3V 63.20 20.40 16.20 79.40 4.20 1290
Mean 84.66 27.04 20.86 105.52 3.57 2324

 

87

on PC3 axis for Figure 3 (Station I 2).

Appendix 6: Means of characters measured highly loading

 

FRZ FR3 KR1

FR1

SH

Clone

 

721.1 71.38“.l 54041..“4461519539647 90 97910235
.J.J.J.J .O.J.J.J .J.J .J.I.J.4.J.4 .I.4.D.D.J.J.J.J.J.J.J.J.J .J.J.J.J.J.J.J.4

ooooooooooooooo1000000000000000000000000000000

SN6finws awnnwnw7flfinmnwnanMnnflnn7nwn7wanwnn%7nfi

0.mommmmmooooooooooooooooo000000000000000000000

ooooooooooooooooo00000000000000000000000000000

annfla9flwwwfla6 n:awaflvfi xuaawwwwwwfi 5w 9mnn&flfi7

ooooooooooooooooooooooooooooooooooooo000000000

33542 .353JJ553654445222J5522 225422 .Aw3233223214.

V.

JJ UV 3.

. O V JRJ13 02 I
.0 t luRu Ru 141-valla at, e
sra P. 35 [RI// u Rn
EA»! n l/RMZMZttt 3R0 L2.‘
0 ¢.K O .t 0 A67... .:4 II .I U w U 1|.IHN .u./.l
9x..lny M “RRLRRR R L11nmg/la..lto Ohh/l 319”
a.l.m t SZB491R2RRRRLBII uaa—IH.‘ cc..l..ll.S/e°
r s a C r s .Iq‘qCIOAI1I1IRv7_ .3.6.4.‘.6 t.t r 0 a a U.l.l .K.K r r.l I." Z
al.1v-l r W.‘ Pli/l/l/l/I/l/lrrOleOlf hhhtdcn
era“ a rhuzz122211112111aah.leek..l..l 999.19-I..I
FCCBDHTFCI 8888883388BBBBHHKTKKSTvI-ITT IIlAAIT

0.60

0.74

0.50

0.68

0.39

Mean

 

88

Appendix 7: Means of characters measured highly loading
on Pol axis for Figure 4 (station I 3).

 

 

Clone NWT NW NTH KWT KW NVOL KVOL
Tuono 3.24 24.90 16.94 1.22 15.10 13592 2732
Ferragnes 4.62 23.32 17.06 1.76 14.46 15099 3920
82/19BL 2.52 18.28 14.02 0.70 10.24 8535 1432
B1/13R 3.06 21.28 13.34 1.10 12.46 9839 2222
B1/15R. 2.72 20.70 14.88 0.94 12.40 9940 2093
Bl/4R. 3.72 21.56 15.64 1.20 12.22 11931 2537
81/16R 2.60 19.80 13.08 0.70 11.66 7132 1449
Bl/17R, 2.38 22.32 14.80 0.90 11.72 1135 1877
BZ/2R 1.26 16.06 12.60 0.68 9.54 5535 1314
Bl/2R 4.16 22.74 14.24 1.16 13.36 11095 2345
Bl/lL 2.12 19.18 13.16 0.86 11.94 9402 1840
Bl/7R 1.98 19.24 15.64 0.92 12.64 8758 2001
82/7R 3.26 21.86 15.22 1.48 13.44 12414 3142
Bl/4L 3.48 19.14 13.88 1.22 11.76 10441 2327
Hart/18J 2.86 20.64 12.98 0.82 13.40 7516 1826
Khorbat/3J 2.78 17.86 13.50 0.84 11.48 7210 1885
Khorbat/6J 2.84 18.60 14.02 0.86 10.90 7503 1716
Kelaa/7R 5.12 28.36 20.12 1.52 16.54 18906 3464
Amek./1J 3.10 19.88 15.34 0.96 12.44 8416 2071
Amek./3J 5.68 25.06 16.56 1.26 14.54 15233 2450
Tiflit/ZV 2.36 19.76 14.80 0.78 10.52 8558 1694
Tiflit/ZR 1.90 16.62 12.12 0.56 10.84 4656 1293
Toundout/8J 1.10 16.30 10.92 0.78 10.40 4961 1702
Toundout/3J 2.66 19.12 13.56 0.92 12.64 7938 2210
Toundout/lR 3.74 21.46 14.04 0.94 13.34 9490 2030
Tiliwine/8V’ 2.64 22.02 14.84 1.42 13.60 11538 2988
Tiliw./8TER. 5.36 23.36 16.96 1.36 15.00 16151 3015
Ighri/lZB 4.84 22.28 15.96 1.46 14.22 11447 3144
Ighri/lBB 5.10 24.44 16.42 1.60 15.76 14402 3532
Ait Sao./2V’ 2.04 18.26 13.76 1.08 11.82 8204 2396
Ait Sao./4V’ 3.48 22.40 15.20 1.32 12.98 11858 2681
Ait Sao./5V’ 4.82 24.32 19.58 1.48 14.36 15269 3401
Ait Sao./6V 2.10 18.52 13.86 1.00 11.26 7482 2071
Ait Sao./S3 2.54 17.70 13.98 0.64 11.22 6540 1444
‘Agdz/IBL 2.78 19.34 13.04 0.84 12.22 7578 1863
Ircheg/ZR. 3.58 21.50 15.74 0.92 13.14 -9752 1899
Tamkas./3R. 3.46 22.84 16.02 1.14 12.82 10903 2462
Tinz./5R 3.64 23.26 16.46 0.96 12.86 11597 2131
Tinz./3V 2.76 22.96 15.26 0.96 15.62 9573 2534
Mean 3.19 20.95 14.86 1.06 12.74 10193 2285

 

89

 

 

Appendix 8: Means of characters measured highly loading
on P02 axis for Figure 4 (station I 3).

 

 

Clone NR1 NR2 KR1 KR2 NLz KL‘ SNBRE
Tuono 0.77 0.52 0.67 0.36 32.16 22.32 6.2
Ferragnes 0.61 0.45 0.50 0.32 37.82 28.86 7.0
82/19BL 0.55 0.42 0.45 0.27 33.18 22.52 7.0
Bl/13R 0.61 0.38 0.49 0.28 34.54 25.12 8.4
Bl/15R 0.64 0.46 0.54 0.32 32.60 22.80 7.0
Bl/4R 0.61 0.44 0.47 0.31 35.32 25.56 6.4
Bl/6R 0.72 0.47 0.58 0.31 27.50 19.86 6.8
Bl/17R 0.66 0.43 0.49 0.28 33.84 23.64 7.4
82/2R 0.59 0.46 0.50 0.38 27.32 19.00 6.8
Bl/2R 0.66 0.41 0.50 0.24 34.12 26.54 7.4
Bl/lL 0.51 0.35 0.47 0.23 37.16 25.08 7.4
B1/7R 0.67 0.54 0.62 0.38 28.60 20.32 7.8
BZ/7R 0.58 0.40 0.51 0.34 37.34 26.12 7.8
Bl/4L 0.48 0.35 0.43 0.26 38.22 27.12 8.6
Hart/18J 0.73 0.46 0.65 0.32 27.94 20.58 7.8
Khorbat/3J 0.60 0.45 0.55 0.38 29.60 20.48 7.0
Khorbat/6J 0.65 0.49 0.54 0.39 28.40 20.10 7.6
Kelaa/7R 0.85 0.60 0.67 0.34 33.04 24.46 7.2
Amek./1J 0.72 0.56 0.62 0.41 27.40 20.08 7.4
Amek./3J 0.68 0.45 0.69 0.36 36.66 21.94 7.6
Tiflit/ZV 0.67 0.50 0.48 0.34 29.20 21.54 7.8
.Tiflit/ZR 0.72 0.52 0.62 0.40 23.06 17.26 8.2
Toundout/8J 0.58 0.39 0.48 0.36. 27.78 21.30 7.2
Toundout/BJ 0.62 0.44 0.53 0.32 30.52 23.46 7.4
Toundout/lR 0.68 0.44 0.60 0.30 31.38 22.18 8.0
Tiliw./8V 0.63 0.42 0.54 0.32 34.92 24.88 6.2
Tiliw./8TER. 0.57 0.41. 0.56 0.28 40.88 26.50 7.2
Ighri/lZB 0.69 0.49 0.56 0.351 32.10 25.00 8.2
Ighri/13B 0.70 0.47 0.63 0.34 34.64 24.96 8.2
Ait Sao./2V 0.56 0.42 0.48 0.33 32.50 24.34 7.8
Ait Sao./4V 0.64 0.43 0.52 0.33 34.80 24.76 8.6
Ait Sao./5V 0.76 0.61 0.60 0.41. 31.84 23.92 ‘7.2
Ait Sao./6V 0.63 0.47 0.50 0.36 29.08 22.52 6.6
Ait Sao./S3 0.67 0.53 0.58 0.34 26.34 19.34 6.6
Agdz/lBL 0.64 0.43 0.56 0.32 29.92 21.80 7.4
Ircheg/ZR 0.74 0.54 0.65 0.36 28.78 20.00 6.8
Tamkas./3R. 0.76 0.54 0.58 0.40 29.70 21.88 8.2
Tinz./5R 0.77 0.54 0.65 0.42 30.20 19.72 7.8
Tinz./3V 0.85 0.56: 0.77 0.38 26.74 20.16 7.0
Mean 0.67 0.47 0.56 0.34 31.73 22.77 7.5

 

‘Characters loading negatively on P02.

90

 

Appendix 9: Means of characters measured highly loading
on PC3 axis for Figure 4 (station # 3).

 

 

Clone LA‘I'z TOTSPRS 82Lz Sle SPRS NODE‘
Tuono 0.2 7.0 15.0 17.2 0.49 1.4
Ferragnes 0.0 6.6 20.8 11.6 0.33 0.0
B2/193L 0.0 0.0 14.3 10.8 0.00 0.0
B1/13R 0.0 2.8 14.9 21.2 0.19 0.0
Bl/15R 0.0 5.6 10.5 11.4 0.53 0.0
B1/4R 0.0 3.8 11.1 9.7 0.33 0.0
Bl/16R 0.0 3.4 12.9 7.9 0.27 0.0
B1/17R 0.4 0.0 9.4 6.8 0.00 1.8
BZ/ZR 0.0 6.8 13.7 22.5 0.51 0.0
Bl/2R 0.8 3.0 11.4 7.7 0.30 1.8
Bl/lL 0.8 0.8 10.2 13.8 0.09 1.8
Bl/7R 0.0 4.4 14.7 8.3 0.30 0.0
B2/7R 0.0 2.8 11.0 13.7 0.27 0.0
Bl/4L 0.0 1.4 15.7 9.0 0.08 0.0
Hart/18J 0.0 6.0 17.0 10.2 0.36 0.0
Khorbat/3J 0.0 3.2 9.4 20.8 0.37 0.0
Khorbat/6J 1.2 0.8 36.8 41.2 0.03 6.0
Kelaa/7R 2.0 3.4 14.7 38.1 0.28 1.4
Amek./1J 0.6 4.6 22.2 22.6 0.21 2.6
Amek./3J 1.4 1.6 20.6 19.2 0.12 5.4
Tiflit/ZV 0.0 4.6 10.7 7.2 0.47 0.0
Tiflit/ZR 0.0 6.6 17.9 7.1 0.38 0.0
Toundout/8J 0.2 2.4 13.6 19.3 0.15 1.6
Toundout/BJ 0.0 2.4 8.8 10.4 0.31 0.0
Toundout/1R 0.0 4.8 11.8 11.2 0.43 0.0
Tiliwine/BV 0.0 2.6 11.0 10.4 0.28 0.0
Tiliw./8TER 0.2 2.0 12.1 15.2 0.18 1.6
Ighri/lZB 0.0 4.6 10.2 16.6 0.45 0.0
Ighri/13B 0.4 2.6 9.9 13.2 0.26 1.8
Ait Sao./2V 0.0 5.2 8.4 8.8 0.66 0.0
Ait Sao./4V 0.0 0.4 12.1 21.1 0.05 0.0
Ait Sao./5V 0.0 1.8 13.0 17.2 0.10 0.0
Ait Sao./6V 0.0 3.6 8.9 8.3 0.40 0.0
Ait Sao./83 0.0 6.2 10.0 5.2 0.63 0.0
Agdz/lBL 0.0 4.4 9.2 7.4 0.46 0.0
Ircheg/ZR 0.4 2.6 13.6 14.0 0.20 1.8
Tamkas./3R 0.0 4.2 11.8 16.9 0.34 0.0
Tinz./5R 0.0 7.4 16.5 17.0 0.44 0.0
Tinz./3V 0.0 6.2 12.3 8.1 0.51 0.0
Mean 0.22 3.66 13.5 14.3 0.30 0.97

 

zCharacters loading negatively on PC3.

91

Appendix 10: Eigenvalues of the corrrelation matrix for the
seven first PC's at station #1.

 

 

PC axis eigenvalue difference proportion cumulative
PC1 8.435 2.343 0.324 0.324
PC2 6.083 2.515 0.234 0.558
PC3 3.567 1.245 0.137 0.695
PC4 2.323 0.577 0.089 0.785
PCS 1.746 0.220 0.067 0.852
PC6 1.526 0.676 0.059 0.910
PC7 0.845 0.300 0.033 0.943

 

92

 

Appendix 11: Eigenvalues of the corrrelation matrix for the
seven first PC's at station #2.

 

 

PC axis eigenvalue difference proportion cumulative
PC1 6.980 2.745 0.269 0.269
PC2 4.235 0.308 0.163 0.431
PC3 3.927 1.510 0.151 0.582
PC4 2.418 0.365 0.093 0.675
PCS 2.053 0.161 0.079 0.754
PC6 1.892 0.816 0.073 0.827
PC7 1.077 0.093 0.869

0.041

 

93

Appendix 12: Eigenvalues of the corrrelation matrix for the
seven first PC's at Station #3.

 

 

PC axis eigenvalue difference proportion cumulative
PC1 8.649 2.309 0.254 0.254
PC2 6.340 2.323 0.187 0.441
PC3 4.017 0.835 0.118 0.559
PC4 3.182 0.515 0.094 0.653
PCS 2.667 0.784 0.078 0.731
PC6 1.883 0.588 0.055 0.786
PC7 1.296 0.034 0.038 0.825

 

94

Appendix 13: Almond nuts and kernels of 13 Moroccan
selections.

 

 

95

GENERAL CONCLUSION :

Sour cherry Prunus c rasus, commonly known as a self-
compatible Prunus species, exhibited gametophytic self-
incompatibility by pollen tube growth inhibition in the style.
Pollen tube growth and quantity of pollen tubes reaching the
ovary suggested the presence of different levels of
incompatibility reaction, thus indicating the existence of
self-incompatible, partially self-incompatible, and self-
compatible cultivars.

Self-incompatibility in Prunus dulcis resulted in the
actual genetic variability that exists in different almond
producing countries. However, the extensive use of few
superior genotypes in breeding programs to improve almond
production may reduce the available genetic diversity, and
restrict future genetic gain. 'Nonpareil' and 'Mission'
cultivars, representing respectively 65% and 25% of commercial
almond cultivars in California, are still used extensively as
gene sources in California breeding programs. Russian breeders
use 'Nikitski 62' cultivar as the major genotype in their
breeding programs. In Western Europe and North Africa, Italian
cultivars 'Tuono' and 'Cristomorto' , and French cultivars 'Ai'
and 'Ferragnes' are being extensively used for almond cultivar
improvement. . This has resulted in increased inbreeding in
almond cultivars, particularly in the United States, as well

as in varying degrees of coancestry relationships among almond

96

cultivars in the U.S, Russia, Israel, France and Spain.
Germplasm exchange and the use of the same progenitors has led
to coancestry relationships among U.S, Russian, and Israeli
almond cultivars, and between French and Spanish cultivars.
In North Africa, and particularly in Morocco, almond
breeding programs are at their beginning, and almonds are
propagated mostly by seedlings, and grown under different
environments. As a consequence, genetic diversity may be
present and almond ecotypes may have developed. Morphological
studies of selected clones of almonds from Southern Morocco
and introduced cultivars revealed that Moroccan selections are
characterized by small leaves, large fruit variability, and
limited fruiting potential. However, some superior Moroccan
genotypes were identified. IntrOduced cultivars had long and
wide leaves, and.good ratio of shoot growth.: yield.potential.
Even though some Moroccan genotypes from the same area of
origin clustered together, there was no evidence of separate
ecotypes existing in southern Morocco. The morphological
variation observed among genotypes indicates the existence of
genetic potential for the development of breeding programs to

improve almond production in Morocco.

97

 

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