Performance and genotypic variability in diverse date palm (Phoenix dactylifera L.) cultivars for fruit characteristics

Abstract

Identification of date palm cultivars is primarily based on fruit characteristics while the morphological characteristics of the trees have less importance. Exploring genotypic variability in date palm cultivars is indispensable for enhancing sustainable utilization, dynamic conservation, and the development of new germplasm. Accordingly, 16 diverse date palm cultivars from different sites in Saudi Arabia were evaluated for various fruit characteristics in the present study. The assessed date palm cultivars exhibited highly significant differences in all studied fruit characteristics. The highest fruit length was displayed by Um-Kbar, Wannana, Medjool, and Sefri cultivars, while the largest width of fruits was assigned to Rushodia, Um-Kbar, and Wannana. Fruit weight and volume were significantly higher in Um-Kbar, and Medjool than those in other cultivars. On the other hand, the Medjool, Hilali, Thawee, and Sefri cultivars recorded the highest soluble solid content. Um-Kbar, Khesab, and Hilali recorded the highest acidity percentage (more than 0.60%). The phenotypic variance (PV) values of all studied characteristics were higher than the genotypic variances (GV). Additionally, the phenotypic coefficient of variation (PCV) was larger than the genotypic coefficient of variation (GCV). The highest PCV and GCV values were observed for the length of basal appendages, whereas the lowest values were assigned for fruit length. All studied characteristics displayed high heritability (h²b), varying from 99.61 (of fruit volume) to 84.49 (of seed length). Expected genetic advance (GA) values ranged from high for fruit weight to low for seed width. The results of the principal component analysis revealed that most of the variation was explained by the two components. Accordingly, they were employed to construct PC biplot to present the association between evaluated cultivars and studied fruit characteristics. The studied cultivars were clustered based on fruit characteristics into six groups ranging from high to low performance.

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https://doi.org/10.1007/s10722-023-01719-0
RESEARCH ARTICLE
Performance andgenotypic variability indiverse date palm
(Phoenix dactylifera L.) cultivars forfruit characteristics
MesferM.Alqahtani· MaysounM.Saleh· KhairiahM.Alwutayd·
FatmahA.Safhi· SalahA.Okasha · MohamedA.Abdelsatar·
MohamedS.M.Ali· MagdiI.Saif· AmiraA.Ibrahim· KhaledF.M.Salem
Received: 10 July 2023 / Accepted: 20 August 2023
© The Author(s), under exclusive licence to Springer Nature B.V. 2023, corrected publication [2023]
Abstract Identification of date palm cultivars is pri-
marily based on fruit characteristics while the mor-
phological characteristics of the trees have less impor-
tance. Exploring genotypic variability in date palm
cultivars is indispensable for enhancing sustainable
utilization, dynamic conservation, and the develop-
ment of new germplasm. Accordingly, 16 diverse date
palm cultivars from different sites in Saudi Arabia
were evaluated for various fruit characteristics in the
present study. The assessed date palm cultivars exhib-
ited highly significant differences in all studied fruit
characteristics. The highest fruit length was displayed
by Um-Kbar, Wannana, Medjool, and Sefri culti-
vars, while the largest width of fruits was assigned to
Rushodia, Um-Kbar, and Wannana. Fruit weight and
volume were significantly higher in Um-Kbar, and
Medjool than those in other cultivars. On the other
hand, the Medjool, Hilali, Thawee, and Sefri culti-
vars recorded the highest soluble solid content. Um-
Kbar, Khesab, and Hilali recorded the highest acidity
percentage (more than 0.60%). The phenotypic vari-
ance (PV) values of all studied characteristics were
Supplementary Information The online version
contains supplementary material available at https:// doi.
org/ 10. 1007/ s10722- 023- 01719-0.
M.M.Alqahtani
Department ofBiological Sciences, Faculty ofScience
andHumanities, Shaqra University, Ad-Dawadimi11911,
SaudiArabia
M.M.Saleh
Genetic Resources Department, General Commission
forScientific Agricultural Research, Damascus, Syria
K.M.Alwutayd· F.A.Safhi
Department ofBiology, College ofScience, Princess
Nourah bint Abdulrahman University, P.O. Box84428,
Riyadh11671, SaudiArabia
S.A.Okasha(*)
Department ofAgronomy, Faculty ofAgriculture, Suez
Canal University, Ismailia41522, Egypt
e-mail: salah.okasha@agr.suez.edu.eg
M.A.Abdelsatar
Oil Crops Research Department, Field Crops Research
Institute, Agricultural Research Center, Giza, Egypt
M.S.M.Ali· M.I.Saif
Department ofHorticulture, Faculty ofAgriculture, Suez
Canal University, Ismailia41522, Egypt
A.A.Ibrahim
Botany andMicrobiology Department, Faculty ofScience,
Arish University, Al-Arish45511, Egypt
K.F.M.Salem
Department ofBiology, College ofScience
andHumanities, Shaqra University, Qwaieah, SaudiArabia
K.F.M.Salem
Plant Biotechnology Department, Genetic Engineering
andBiotechnology Research Institute (GEBRI), Menoufia
University, P.O. Box, 79, SadatCity, Egypt
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higher than the genotypic variances (GV). Addition-
ally, the phenotypic coefficient of variation (PCV)
was larger than the genotypic coefficient of varia-
tion (GCV). The highest PCV and GCV values were
observed for the length of basal appendages, whereas
the lowest values were assigned for fruit length. All
studied characteristics displayed high heritability
(h2b), varying from 99.61 (of fruit volume) to 84.49
(of seed length). Expected genetic advance (GA) val-
ues ranged from high for fruit weight to low for seed
width. The results of the principal component analy-
sis revealed that most of the variation was explained
by the two components. Accordingly, they were
employed to construct PC biplot to present the asso-
ciation between evaluated cultivars and studied fruit
characteristics. The studied cultivars were clustered
based on fruit characteristics into six groups ranging
from high to low performance.
Keywords Mean performance· Genotypic and
phenotypic coefficient of variation· Heritability·
Genetic advance· Principal component analysis·
Cluster analysis
Introduction
Date palm (Phoenix dactylifera L.) is one of the
most economical fruit crops (Harkat et al. 2022). It
contributes to environmental conservation, agricul-
tural sustainability, and food security worldwide in
arid regions with limited water resources (Haj-Amor
et al. 2020; Dhaouadi etal. 2021). Its global culti-
vated area in 2021 was 1.30million hectares, produc-
ing 9.66 million tons (FAOSTAT 2023). Date palm
is an extensively grown fruit tree in the predominat-
ing climate of long, dry summers of the Middle East
and North Africa region (El Hadrami and Al-Khayri
2012). Iraq, Algeria, Iran, Saudi Arabia, Pakistan,
Tunisia, Morocco, Egypt, and the United Arab Emir-
ates represent the highest cultivated areas of date
palm, respectively (FAOSTAT 2023). Consequently,
Egypt, Saudi Arabia, Iran, and Algeria displayed the
highest production in 2021 with 1.75, 1.57, 1.30, and
1.19million tons, presenting 18.10, 16.22, 13.50, and
12.31% of the global production in the same order
(FAOSTAT 2023). Date fruit is a rich energy source,
with a high content of carbohydrates and essential
minerals (Fe, Cu, S, K, and Mg), and vitamins as well
as thiamine, riboflavin, and folic acid. Accordingly,
date fruits are considered a vital industrial resource
worldwide (Mrabet etal. 2016).
Investigating genetic diversity becomes irreplace-
able before planning a proper breeding strategy for
improving different crops, particularly under current
abrupt climate changes (Ponce-Molina et al. 2012;
Kamara etal. 2021b; Müller etal. 2021; Broccanello
et al. 2023). Exploring genetic variability allows
breeders to identify promising germplasm with differ-
ent characteristics to develop desired genotypes under
different environmental conditions (Gracia et al.
2012; Abaza et al. 2020; Gayacharan et al. 2022).
Besides, considerations of genetic variability can
assist breeders in recognizing gene pool variations
and speed up the selection of preferred character-
istics and resilience to climate change (Gharib etal.
2020; Habibullah et al. 2021; Kamara et al. 2021a;
Ali etal. 2023). Thus, examining genetic variability
in the available germplasm is valuable for enhancing
genetic diversity and developing improved genotypes
(ElShamey etal. 2022; Misganaw etal. 2023).
Date palm is a perennial monocotyledonous,
angiosperm, diploid (2n = 36), and dioecious plant
belonging to the Arecaceae family (Asaf etal. 2021).
Its female and male flowers are borne on separate
trees. Some cultivars are planted with seeds. Accord-
ingly, a large diversity is detected among their traits.
The wild and seedy date palm species are morpho-
logically similar to the domesticated form (Gros-
Balthazard et al. 2016). Certain date palm cultivars
have narrow morphological and fruit characteristics.
This situation requires evidence to prove phylogenetic
relationships at the interspecific level, highlighting
the significance of estimating the genotypic diversity
among date palm cultivars. Moreover, the success of
breeding programs depends on exploring the quantity
and distribution of genotypic variability in the exist-
ing genetic pool (Khierallah etal. 2017). Therefore,
assessing phenotypic and genotypic variability in date
palm cultivars is decisive in determining the geno-
typic potential for breeding efficiency (Ahmed etal.
2021). This supports sustainable utilization, dynamic
conservation, phylogenetic relationships, and the
development of new germplasm (Elsafy et al. 2016;
Saboori et al. 2020). In this context, numerous pre-
vious reports demonstrated genetic variability in date
palm (Salem etal. 2008; El Kadri etal. 2019; Ahmed
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et al. 2021; Ahmad et al. 2023; Elboghdady et al.
2023).
Climate change substantially harms agricultural
production (Shabani etal. 2014; Allbed etal. 2017).
Local adaptive genetic variations can support the
potential of date palms to cope with abrupt environ-
mental changes (Naqvi etal. 2015; El Kadri et al.
2019). Therefore, exploring genotypic variations of
local genotypes is essential, in particular under the
current climate change. Keeping in view the signifi-
cance of assessing genotypic variability in local date
palms genotypes and the lack of available informa-
tion about that in the Middle East region, the present
study aimed at (i) estimating phenotypic and geno-
typic variability among diverse sixteen date palm
cultivars collected from different regions, (ii) deter-
mining the degree of diversity of fruit characteris-
tics using multivariate analysis, and (iii) defining the
contribution of fruit characteristics to the total vari-
ation and identifying their heritability and genetic
advance.
Materials andmethods
Plant materials
Sixteen local date palm (Phoenix dactylifera L.,
2n = 36) cultivars from different sites in Saudi
Arabia (Table 1) were evaluated for various fruit
characteristics. The assessed cultivars are presented
in Table2.
Data collection
Date palm fruits from 16 cultivars were harvested in
the first week of November 2021 from different sites
Table 1 Collection sites of
studied date palm cultivars
in Saudi Arabia
Genotype Region Latitude Longitude
Segae Riyadh 24.42° N 46.71° E
Sukkari Al-Qassim 25.81° N 42.87° E
Thawee AL Madinah AL Munawwarah 24.46° N 39.61° E
Sukkari Ahmr Al-Qassim 25.81° N 42.87° E
Rushodia Al-Qassim 25.81° N 42.87° E
Ruthana AL-Madinah Al-Munawwarah 24.46° N 39.61° E
Nabtat Ali Al-Qassim 25.81° N 42.87° E
Shaishee Al-Ahsa 25.28° N 49.48° E
Wannana Al-Qassim 25.81° N 42.87° E
Hilali Al-Ahsa 25.28° N 49.48° E
Dekhaini Al-Qassim 25.81° N 42.87° E
Khalas Al-Ahsa 25.28° N 49.48° E
Sefri Bisha district 20.11° N 42.62° E
Medjool Al-Qassim 25.81° N 42.87° E
Khesab Al-Ahsa 25.28° N 49.48° E
Um-Kbar Al-Qassim 25.81° N 42.87° E
Table 2 The 16 date palm cultivars were assessed in this study
No Cultivars name Tamar stage color Maturity class
1 Segae Reddish-brown Mid-season
2 Sukkari Brown Mid-season
3 Thawee Dark-red Mid-season
4 Sukkari Ahmr Brown Mid-season
5 Rushodia Brown Mid-season
6 Ruthana Yellowish-brown Early
7 Nabtat Ali Maroon Mid-season
8 Shaishee Reddish-brown Mid-season
9 Wannana Dark-brown Mid-season
10 Hilali Brown Late
11 Dekhaini Dark-brown Early
12 Khalas Brown Mid-season
13 Sefri Brown Mid-season
14 Medjool Brown Mid-season
15 Khesab Dark Late
16 Um-Kbar Brown Late
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in Saudi Arabia. One kilogram of each cultivar was
collected for evaluating fruit characteristics. The
fruits were washed with tap water and dried by van
air for 2h. The recorded fruit characteristics are pre-
sented in Table3. Fruit and seed length and width
were measured in cm by Vernier caliper. Fruit and
seed weight and flesh weight were measured in gm
by electric balance. Fruit and seed volume (cm3) was
determined according to Horwitz and Latimer (2005).
Fruit flesh percentage was calculated from the per-
centage of flesh to the whole fruit weight. The soluble
solid content was measured using an Abbe refractom-
eter Model DR-A1-Plus, Tokyo, Japan. The acidity
percentage was calculated as malic acid by measuring
the titration of 0.1N of NaOH, equivalent per 100g
FW.
Statistical analysis
The data collected were analyzed using one-way
ANOVA. Tukey’s HSD test was performed to evalu-
ate statistical significance among the cultivars. The
datasets were first tested with normality tests and
then subjected to variance analysis. The 16 culti-
vars were divided into groups using two methods:
unweighted pair group system of average linkage
(UPGMA) and principal component analysis. The
experimental results were statistically analyzed
using R statistical software version 4.1.1 for multi-
variate analysis.
Results
Performance of date palm cultivars
The assessed cultivars exhibited highly signifi-
cant differences in all assessed fruit characteris-
tics (Table 4). The longest fruits were recorded by
Um-Kbar, Wannana, Medjool, and Sefri (more than
4.5 cm), whereas Ruthana had the shortest ones
(3.38 cm). Rushodia and Um-Kbar had the largest
fruit width (more than 3.3 cm), which was signifi-
cantly high compared to the other cultivars, and Hilali
had the lowest diameter (less than 2.0cm). The fruit
weight was higher in Um-Kbar, Medjool, and Wan-
nana (more than 15 g) and lower in Ruthana and
Hilali (less than 10g). Fruit volume was significantly
higher in Um-Kbar and Medjool (more than 15 cm3)
than that in other cultivars. Fruit weight, dimen-
sions, and volume indicated that Um-Kbar, Wannana,
and Medjool had the largest size in their fruits, sig-
nificantly different from Ruthana and Hilali (small
fruits). The flesh weight was significantly higher in
Um-Kbar and Medjool (more than 17 g) compared
to Hilali and Dekhaini, which had the lightest weight
of fruit flesh (less than 9g). Medjool, Hilali, Thawee,
Sukkari, and Sefri exhibited the highest soluble solid
content (SSC, more than 50%). On the other hand,
Khesab recorded the lowest value (39.45%) compared
to the other cultivars (Table 5). Um-Kbar, Khesab,
and Hilali displayed the highest acidity percentage
(more than 0.60%). Otherwise, Khalas, Dekhaini,
Ruthana, and Segae had the lowest acidity percent-
age (less than 0.50%). Um-Kbar, Dekhaini, Sefri,
Shaishee, and Sukkari Ahmr displayed the highest
values of seed length, seed weight, and seed width,
whereas Ruthana, Khesab, and Thawee possessed the
lowest ones.The assessed cultivars exhibited highly
significant differences in all assessed fruit charac-
teristics (Table4). The longest fruits were recorded
by Um-Kbar, Wannana, Medjool, and Sefri (more
than 4.5cm), whereas Ruthana had the shortest ones
(3.38 cm). Rushodia and Um-Kbar had the largest
fruit width (more than 3.3 cm), which was signifi-
cantly high compared to the other cultivars, and Hilali
had the lowest diameter (less than 2.0cm). The fruit
Table 3 Measured fruit traits in 16 date palm cultivars
Trait Unit Abbreviation
Fruit weight g FEW
Fruit length cm FL
Fruit width cm FWI
Fruit diameter cm FD
Fruit volume cm3FV
Flesh weight g FLW
Fruit flesh % FF
Length of basal appendages mm LBA
Seed weight g SW
Seed length cm SL
Seed width cm SWI
Seed diameter cm SD
Seed volume cm3SV
Soluble solid content % SSC
Acidity percentage % AP
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Table 4 Performance for the fruit traits of the assessed 16 date palm cultivars
Mean values followed by the same letter in each column show nonsignificant differences at the P < 0.05 probability level
Cultivar Fruit weight (g) Fruit length (cm) Fruit width (cm) Fruit diameter(cm) Fruit volume (cm3) Flesh weight (g) Fruit flesh (%)
Segae 13.79e4.10cd 2.20d8.14bc 12.03e12.62d91.47b
Sukkari 13.12f3.54ef 2.69b7.90 bc 10.03f11.89e90.58bcd
Thawee 10.63i3.94d2.17d3.33f9.83f9.92h93.29a
Sukkari
Ahmr
11.65h3.69e2.53bc 8.06bc 8.87gh 10.45g89.68d
Rushodia 14.71d4.08cd 3.43a5.43e13.67cd 13.46c91.53b
Ruthana 9.87j3.38f2.53bc 8.02bc 8.33h8.94i90.54bcd
Nabtat
Ali
12.77fg 3.54ef 2.52bc 7.92bc 13.37d11.43ef 89.57d
Shaishee 12.20g3.66ef 2.33d7.50d9.00gh 11.13f91.26b
Wannana 15.23c4.63b2.71b8.86bc 14.17c13.90c91.29b
Hilali 9.60j4.27c1.76f9.53ab 8.33h8.48i88.27e
Dekhaini 10.14ij 4.08cd 2.01e7.70bc 8.27h8.70yi85.79f
Khalas 10.71i3.61ef 2.19d6.93d7.60i9.77h91.25b
Sefri 12.47g4.54b2.28d7.17d8.67gh 11.33f90.85bcd
Medjool 18.42b4.57b2.69b7.03d15.17b17.25b93.66a
Khesab 10.55i3.62ef 2.37cd 7.06d9.17g9.61h91.11bc
Um-Kbar 23.07a5.04a3.34a10.53a18.17a20.73a89.84cd
P-value < 0.001 < 0.001 < 0.001 < 0.001 < 0.001 < 0.001 < 0.001
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weight was higher in Um-Kbar, Medjool, and Wan-
nana (more than 15 g) and lower in Ruthana and
Hilali (less than 10g). Fruit volume was significantly
higher in Um-Kbar and Medjool (more than 15 cm3)
than that in other cultivars. Fruit weight, dimen-
sions, and volume indicated that Um-Kbar, Wannana,
and Medjool had the largest size in their fruits, sig-
nificantly different from Ruthana and Hilali (small
fruits). The flesh weight was significantly higher in
Um-Kbar and Medjool (more than 17 g) compared
to Hilali and Dekhaini, which had the lightest weight
of fruit flesh (less than 9g). Medjool, Hilali, Thawee,
Sukkari, and Sefri exhibited the highest soluble solid
content (SSC, more than 50%). On the other hand,
Khesab recorded the lowest value (39.45%) compared
to the other cultivars (Table 5). Um-Kbar, Khesab,
and Hilali displayed the highest acidity percentage
(more than 0.60%). Otherwise, Khalas, Dekhaini,
Ruthana, and Segae had the lowest acidity percent-
age (less than 0.50%). Um-Kbar, Dekhaini, Sefri,
Shaishee, and Sukkari Ahmr displayed the highest
values of seed length, seed weight, and seed width,
whereas Ruthana, Khesab, and Thawee possessed the
lowest ones.
Variability, heritability, and genetic advance
The results of genetic parameters were presented in
Table6 for all studied characteristics. The phenotypic
variance values (PV) were higher than the genotypic
variances (GV) for all variables. The phenotypic vari-
ance (PV) ranged from 23.32 (SSC) to 0.01 (acid-
ity percentage), while GV values ranged from 20.53
(SSC) to 0.01 (acidity percentage). Also, the pheno-
typic coefficient of variation (PCV) was higher than
the genotypic coefficient of variation (GCV). The
high values of PCV and GCV were recorded for the
length of basal appendages, whereas the lowest values
were recorded for fruit flesh. Heritability in a broad
sense (h2b) was high for all characteristics and ranged
from 99.28 (fruit weight) to 86.97 (seed volume).
The highest genetic advance (GA) was assigned for
SSC, whereas the lowest one was displayed by acid-
ity percentage. The highest genetic advance per mean
Table 5 Performance for the seed traits of the assessed 16 date palm cultivars
Mean values followed by the same letter in each column show nonsignificant differences at the P < 0.05 probability level
Cultivar Seed length
(cm)
Seed
weight (g)
Seed width
(cm)
Length
of basal
append-
ages (mm)
Seed diameter
(cm)
Seed
volume
(cm3)
Soluble solid
content (%)
Acidity
percentage
(%)
Segae 2.30bc 1.18cde 0.80cd 0.53f2.50cde 1.00ef 52.95bc 0.41c
Sukkari 1.97d1.24cde 0.97bc 0.83f2.93bc 1.47cde 50.68bcd 0.56ab
Thawee 1.95d0.71g0.50fg0.47f2.00f1.64bcd 53.67ab 0.52b
Sukkari Ahmr 2.08cd 1.20cde 0.86bcd 1.91cd 2.19ef 1.20de 50.09bcd 0.55ab
Rushodia 2.00d1.25cde 0.47g0.50f1.98f1.69bcd 44.58ef 0.55ab
Ruthana 1.80d0.93f0.82cd 1.75cde 3.04b0.60f42.09fg 0.41c
Nabtat Ali 2.00d1.33bc 1.00b1.37e3.00b1.87bc 50.64bcd 0.55ab
Shaishee 2.50ab 1.07ef 0.89bc 1.50de 2.73bcd 1.03ef 50.53bcd 0.56ab
Wannana 1.83d1.33bcd 0.80cd 2.47ab 2.50cde 1.02ef 51.33bc 0.56ab
Hilali 1.80d1.13de 0.81cd 2.04bc 2.47de 1.03ef 54.91ab 0.60ab
Dekhaini 2.60a1.44b0.92bc 1.52de 3.04b1.05ef 48.19cde 0.41c
Khalas 2.07cd 0.94f0.69de 0.40f2.37def 1.05ef 48.04cde 0.33d
Sefri 2.70a1.14cde 0.90bc 1.80cde 2.50cde 1.05ef 50.33bcd 0.55ab
Medjool 2.53ab 1.17cde 0.61ef 1.38e2.50cde 1.66bcd 57.16a0.54ab
Khesab 1.27e0.94f0.70de 1.51de 2.47de 2.03b39.45g 0.60ab
Um-Kbar 2.72a 2.34a 1.82a 2.55a 3.72a 3.03a45.95de 0.63a
P-value < 0.001 < 0.001 < 0.001 < 0.001 < 0.001 < 0.001 < 0.001 < 0.001
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(GAM) was recorded in the length of basal append-
ages and the lowest was in the fruit flesh.
Principal component analysis
The results of the principal component analysis rep-
resented 12 components, while only the first three
(PC1, PC2, and PC3) were considered variation rep-
resentatives. The first three components displayed
eigenvalues greater than the unity: 6.64, 1.56, and
1.43, respectively (Table7). Otherwise, the remaining
components were suspended since their eigenval-
ues were less than the unity. The first three compo-
nents explained together 80.3% of the total varia-
tion between the tested cultivars (Table7). The first
component, PC1, explained 55.3% of the total vari-
ation, while 13.2% was due to PC2 and 11.9% was
explained by PC3. The studied characteristics were
distributed differently in the first three principal com-
ponents (Table8). Two characteristics (fruit flesh and
SSC) were highly related to PC2, while the remaining
characteristics were associated with PC1 (Fig.1). The
Table 6 Variability,
heritability, and genetic
advance of the studied traits
in the evaluated 16 date
palm cultivars
PV: phenotypic variance;
GV: genotypic variance;
PCV: phenotypic coefficient
of variation; GCV:
genotypic coefficient of
variation; h2b: heritability
in broad sense; GA: genetic
advance; GAM: genetic
advance per mean and CV.
Trait PV GV PCV GCV h2bGA GAM
Fruit weight 12.63 12.54 27.21 27.12 99.28 7.27 55.66
Fruit length 0.24 0.23 12.23 11.90 94.79 0.96 23.88
Fruit width 0.20 0.19 17.82 17.48 96.29 0.88 35.34
Fruit diameter 2.71 2.54 21.77 21.06 93.61 3.18 41.97
Fruit volume 9.76 9.65 28.62 28.46 98.90 6.36 58.30
Flesh weight 10.80 10.72 27.74 27.63 99.25 6.72 56.71
Fruit flesh 3.60 3.29 2.09 2.00 91.40 3.57 3.94
Seed length 0.17 0.15 19.31 18.32 89.98 0.76 35.79
Seed width 0.09 0.09 36.26 35.37 95.19 0.60 71.09
Seed weight 0.13 0.12 29.74 29.05 95.42 0.71 58.45
Length of basal appendages 0.49 0.47 49.96 48.83 95.50 1.38 98.29
Seed diameter 0.21 0.19 17.61 16.66 89.48 0.85 32.46
Seed volume 0.37 0.33 43.62 40.68 86.97 1.10 78.15
Soluble solid content 23.32 20.53 9.77 9.17 88.04 8.76 17.72
Acidity percentage 0.01 0.01 16.56 16.11 94.68 0.17 32.29
Table 7 Eigenvalue, proportion, and cumulative variation of
12 resulting components
Principal
components
Eigenvalue Proportion of
variation %
Cumulative
variation %
PC1 6.641 0.553 0.553
PC2 1.556 0.132 0.685
PC3 1.434 0.119 0.804
PC4 0.940 0.078 0.882
PC5 0.534 0.042 0.924
PC6 0.425 0.035 0.959
PC7 0.245 0.020 0.979
PC8 0.101 0.008 0.987
PC9 0.070 0.006 0.993
PC10 0.043 0.004 0.997
PC11 0.010 0.001 0.999
PC12 0.002 0.000 1.000
Table 8 The first three principal components of studied traits
in 16 date palm cultivars
Trait PC1 PC2 PC3
Fruit weight − 0.358 0.171 − 0.064
Fruit length 0.209 0.200 − 0.513
Fruit width − 0.218 0.527 − 0.025
Fruit diameter − 0.285 − 0.108 0.482
Fruit volume − 0.326 0.299 − 0.139
Flesh weight − 0.358 0.133 0.050
Seed weight − 0.334 − 0.062 − 0.131
Seed length − 0.159 − 0.588 − 0.386
Seed width − 0.354 − 0.198 0.112
Seed diameter − 0.283 − 0.259 0.295
Seed volume − 0.247 0.218 − 0.310
Length of basal
appendages
− 0.249 − 0.178 0.340
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distribution of the date palm cultivars in relation to
the first two principal components revealed important
diversity between these cultivars that can be exploited
in the breeding program (Fig. 1). Um-Kbar cultivar
had the best performance of variation compared to
the remaining cultivars (Figs.1 and 2). On the other
hand, PC biplot and heatmap analyses showed that
the Dekhaini, Hilali, Rushodia, and Khalas cultivars
exhibited the worst performance compared to other
cultivars. The studied characteristics were positively
correlated, especially fruit weight, fruit volume, flesh
weight, fruit flesh, and fruit length as they had a small
angle between their axes.
Cluster analysis
The evaluated fruit characteristics were employed
to classify the assessed sixteen date palm culti-
vars. The cultivars were classified into six groups
by hierarchical clustering (Fig.3). Group A com-
prised one cultivar Um-Kbar, with the uppermost
performance in most fruit characteristics. Group B
included three cultivars, Meghool, Rushodia, and
Wannana, which exhibited high values in most fruit
characteristics. Group C gathered three cultivars,
Serfi, Segae, and Shaishee, which displayed mod-
erately high values of fruit characteristics. Group D
contained three cultivars, Nabtat Ali, Sukkari, and
Sukkari Ahmr, which recorded moderately low val-
ues of fruit characteristics. Group E is composed
of four cultivars: Ruthana, Khalas, Thawee, and
Khesab had low performance of fruit characteris-
tics. Group F consisted of two cultivars, Hilali and
Dekhaini, with the lowest performance of all evalu-
ated fruit characteristics.
Discussion
Exploring genotypic variability is fundamental for
supporting sustainable utilization, vigorous con-
servation, and the development of new germplasm,
Fig. 1 Principal component biplot distribution of 16 date palm cultivars and recorded traits
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Genet Resour Crop Evol
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particularly under current climate change (Bedjaoui
and Benbouza 2020; Salem et al. 2020; Mansour
etal. 2021; Sakran etal. 2022; Waheed etal. 2023).
Identification of date palm cultivars is primarily
based on fruit characteristics while the morphological
characteristics of the trees have less importance. Date
palm cultivars are commonly identified by various
fruit characteristics such as shape, dimension, fruit,
and pulp weight (Ahmed etal. 2011; Al-Khayri etal.
2015; Saboori etal. 2021). In the present study, fruit
characteristics of 16 diverse date palm cultivars from
different sites were tested. The assessed cultivars dis-
played highly considerable differences in all evalu-
ated fruit characteristics. Likewise, (Hammadi etal.
2009); (Salomon-Torres et al. 2017); (Ibourki et al.
2021); (Ahmad et al. 2023) elucidated substantial
differences in all tested fruit characteristics among
date palm cultivars. Additionally, the genotypic vari-
ance and genotypic coefficient of variation proved the
presence of inherent genetic differences among the
assessed date palm cultivars concerning studied fruit
characteristics. The observed divergences of fruit
characteristics demonstrated the genetic potential
of these cultivars to improve date palm germplasm.
Moreover, the results revealed that PV and PCV were
slightly higher than genotypic variance and genotypic
coefficients of variation for studied fruit characteris-
tics. These findings prove that the inherent variations
among the assessed date palm cultivars that remain
unaltered by environmental conditions are beneficial
for breeding (Elsafy etal. 2015; Imani etal. 2021).
Furthermore, observed high broad-sense heritability
for studied fruit characteristics proved the effective-
ness of these characteristics for direct phenotypic
selection. These assumptions were reinforced by the
identified genetic advance of these characteristics to
selection through date palm genotypes. Therefore,
these characteristics are a source of diversity among
date palm cultivars and could be employed to select
diverse parents for a future palm date hybridization
program. The results are consistent with the find-
ings of Haider et al. (2015), who assessed diverse
Fig. 2 Heatmap and hierarchical clustering dividing the assessed date palm cultivars into different clusters based on studied fruit
traits. Red and blue colors reveal low and high values for the corresponding fruit traits, respectively
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Pakistani date palm cultivars from different origins
and elucidated highly divergent among the evaluated
cultivars. Besides, Al-Dakheel etal. (2022) disclosed
considerable genotypic variation in fruit characteris-
tics among 18 date palm varieties evaluated in Ara-
bian Peninsula.
The acceptable fruit characteristics of date
palm should contain fruit weight greater than
6g, pulp weight more than 5g, and length higher
than 3.5cm, as depicted by Manickavasagan etal.
(2012). Accordingly, date palm cultivars with
these fruit characteristics are desired by date palm
producers because consumers have more prefer-
ence for the big size and fruit flesh weight of date
palm fruits. In this respect, fruit dimension, weight,
and volume reflected that the Um-Kbar, Med-
jool, and Wannana cultivars exhibited high fruit
weight (more than 15 g), large fruit width (more
than 2.7cm), high fruit volume (more than 1 cm3),
and high flesh weight (more than 13g). Therefore,
these cultivars are preferred due to their large fruit
and flesh weight. On the other hand, the Medjool,
Hilali, Thawee, Sukkari, and Sefri cultivars dis-
played the highest SSC (more than 50%). Cultivars
with high SSC are preferred as their fruits could
be used as a sweetener for various traditional and
modern foods (Manickavasagan et al. 2012). Um-
Kbar, Khesab, and Hilali displayed the highest acid-
ity percentage (more than 0.60%). In contrast, the
Khalas, Dekhaini, Ruthana, and Segae cultivars had
the lowest acidity percentage (lower than 0.50%).
The low acidity of these cultivars was due to the
maturity earliness of these cultivars compared to
late cultivars of Um-Kbar, Hilali, and Khesab. The
reproductive cycle differs considerably between
date palm cultivars grown in the same or different
regions. This allows for the extended harvest season
with dates suitable for fresh consumption, storage,
and processing.
The principal component analysis is a multi-
variate analysis that provides a complete repre-
sentation of all aspects of the evaluated variables
(Ringnér 2008; Bro and Smilde 2014; El-Sanatawy
et al. 2021; Swailam et al. 2021; ElSayed et al.
ABC
D
E
F
Fig. 3 Dendrogram of the evaluated 16 date palm cultivars based on the studied fruit traits
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Genet Resour Crop Evol
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2022; Hamad etal. 2022; Gunsilius and Schennach
2023; Mansour et al. 2023). The first three prin-
cipal components explained most of the observed
overall variation (80.30%). Fruit length contributed
more to the overall diversity and had comparatively
larger values in the first principal component. Fruit
width and volume, followed by seed volume, con-
tributed significantly to the second principal com-
ponent. Fruit diameter and length of basal append-
ages contributed significantly to the third principal
component. Fruit diameter and length of basal
appendages contributed significantly to the three
principal components. Besides, the PC biplot was
constructed to graphically illustrate the associa-
tion between date palm cultivars and recorded fruit
characteristics. Um-Kbar was the closest to fruit
weight and flesh weight, which are the most impor-
tant trait indices of date palm yield. In contrast,
Thawee was the worst one, followed by Khalas and
Rushodia cultivars compared to other cultivars.
The fruit characteristics were positively associ-
ated, particularly fruit weight, flesh weight, fruit
volume, seed volume, and fruit width, as were seed
weight, fruit diameter, fruit length, seed weight,
seed diameter, length of basal appendages, and
seed length. The principal component analysis is
broadly employed to assess the association among
studied characteristics and assessed genotypes in
several published reports. In this respect, Hzaa and
Al_Amiry (2021) proved that principal component
analysis distributed characteristics in aggregate
form and explored genetic variability among evalu-
ated genotypes. Kadri etal. (2021) also employed
PC analysis to explore the association between
morphological characteristics in diverse Tunisian
date palm cultivars.
Cluster analysis was utilized to explore the diver-
gence of cultivars into different clusters (Megahed
et al. 2022; Omar et al. 2022; Kaya et al. 2023;
Morsi etal. 2023). The results reflected the diver-
sification between the evaluated cultivars based
on the studied fruit characteristics. The cultivars
were classified into six different groups ranging
from high to low performance. Um-Kbar, Meg-
hool, Rushodia, and Wannana were classified with
the highest performance of most studied fruit char-
acteristics compared to Ruthana, Khalas, Thawee,
Khesab, Hilali, and Dekhaini, which possessed the
lowest performance. In this context, Salem et al.
(2008) used cluster analysis to identify the diver-
gence among the evaluated genotypes based on
their performance. The intercrossing between wide
diverse genotypes could generate desirable seg-
regants. Similarly, Haider et al. (2015) grouped
Pakistani date palm cultivars from various sources
and observed that there was phenotypic variation
among date palm cultivars, as well as some close
association or heterogeneity within cultivars of
the same provenance. In their study, Kadri et al.
(2021) found that phylogenetic association analysis
revealed that two groups were characterized by a
high level of genetic similarity of inflorescences at
each flowering stage.
Conclusions
The assessed cultivars exhibited considerable phe-
notypic and genotypic heterogeneity or divergence.
Consequently, the date palm improvement program
has much promise through direct selection, crossover,
or hybridization involving scattered clusters, which
can be used to produce viable and possible segregant
populations for later breeding research and mak-
ing potential heterogeneity. The Um-Kbar, Meghool,
Rushodia, and Wannana cultivars displayed the high-
est performance of the most studied fruit characteris-
tics compared to the other evaluated cultivars. Cross-
ing among these cultivars could generate desirable
segregants.
Acknowledgements The authors would like to acknowledge
the Princess Nourah bint Abdulrahman University Research-
ers Supporting Project number (PNURSP2023R318), Princess
Nourah bint Abdulrahman University, Riyadh, Saudi Arabia.
Funding This work was supported by the Princess Nourah
bint Abdulrahman University Researchers Supporting Project
number (PNURSP2023R318), Princess Nourah bint Abdulrah-
man University, Riyadh, Saudi Arabia.
Declarations
Conflict of interest The authors declare no conflicts of inter-
est.
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