Phenotypic Variation of Castanea sativa Mill. Ecotypes in Northern Morocco Based on a Multivariate Analysis of Leaf Morphometrics †
1
Bio-Agrodiversity Team, Laboratory of Applied Botany, Department of Biology, Faculty of Science, Abdelmalek Essaâdi University, Tetouan 93002, Morocco
2
Department of Life Sciences, Polydisciplinary Faculty, Abdelmalek Essaadi University, Larache 92030, Morocco
*
Author to whom correspondence should be addressed.
†
Presented at the 1st International Electronic Conference on Biological Diversity, Ecology and Evolution, 15–31 March 2021; Available online: https://bdee2021.sciforum.net/ .
Biol. Life Sci. Forum 2021, 2(1), 34; https://doi.org/10.3390/BDEE2021-09458
Published: 15 March 2021
(This article belongs to the Proceedings of The 1st International Electronic Conference on Biological Diversity, Ecology and Evolution)
Abstract
:For decades, local and traditional species have been neglected and replaced by industrial and improved species. Sweet chestnut ‘Castanea sativa Mill.’, found in a small area in northern Morocco, is no exception. Indeed, Moroccan ecotypes are neither classified nor characterized. This study aims to evaluate the local genetic resources of Castanea sativa Mill. via a multivariate analysis of the morphometric parameters of its leaves. The study involved 6200 leaves from 31 villages in 3 regions; 10 trees/village and 20 leaves/tree were sampled. Then, eight morphometric parameters were analyzed: lamina length (LL), lamina width (LW), petiole length (PL), distance from the base of the leaf to the widest point of the leaf (DBW), blade area (S), perimeter (P), and ratios LL/LW and LL/DBW. Analysis of the descriptive statistics within and between ecotypes initially showed a large variation in the ten parameters studied. This finding was supported by analysis of variance (ANOVA), which revealed a very highly significant difference (p < 0.0001) for all parameters. Indeed, the analysis of agglomerative hierarchical clustering (AHC) made it possible to group the studied ecotypes into three distinct groups based on the surface. Overall, the high level of variability in leaf morphometric parameters indicates that the region is an important center of genetic diversity for which assessment is crucial for the implementation of conservation and enhancement strategies for this heritage.
1. Introduction
Local and traditional species have been neglected and replaced by industrial and improved species for decades. The chestnut, Castanea sativa Mill., present in a small area in the northwestern part of Morocco, is no exception to this trend. Indeed, Moroccan ecotypes have never been studied.
However, the assessment and characterization of the diversity and ecotype structure are crucial for the implementation of a strategy for the enhancement, conservation, and sustainable use of natural resources [1]. This study fits into this framework and aims, for the first time, to characterize and evaluate the local genetic resources of C. sativa via a morphometric analysis of the leaves. Studies of leaves’ morphological traits have been frequently used by researchers to determine the genetic variability of the chestnut [2,3,4]. Notwithstanding, the chestnut’s blade characteristics, in particular, their size, shape, and anatomy, are largely influenced by developmental, environmental, and cultural factors [5,6]. Nevertheless, many authors confirm that leaf parameters may be appropriate variables for determining the level of genotype variability [2,6,7,8].
2. Materials and Methods
2.1. Study Area
The chestnut leaves were collected in the northwest of Morocco (Figure 1) at the level of 31 villages, divided between three tribal groups (El Smir, Bni Said, and Bni Hozmar), and part of seven watersheds.
The climate of the region is Mediterranean, which is hot and humid in the winter and hot and dry in the summer, with an annual average of 634 mm of precipitation (between 600 mm at low altitude and >1000 mm at high altitude,) and a temperature of 18.3 °C [9]. Generally, the terrain is hilly with a predominance of steep slopes [10].
2.2. Morphometric Parameter
To assess the phenotypic variability of the Moroccan ecotypes, we used an agromorphological characterization of the leaves. The collection of plant material was made from 310 trees distributed among 31 villages, with 10 trees per village. For each chestnut tree, 20 leaves were selected at 2 m in height, making a complete circle around the tree [2,5,8]. The 6200 leaves showed no signs of abnormal growth, mechanical damage, presence of pathogens, or insect infestation. After scanning the leaves, phenotypic variability was investigated by performing a numerical analysis of the images using Image J software (Figure 2). Morphometric measurements took into account the most relevant parameters [2,6,11,12,13]: lamina length (LL), lamina width (LW), distance from lamina base to greatest width (DBW), petiole length (PL), perimeter (P), and blade area (S), and then two ratios were calculated: LL/LW and LL/DBW.
2.3. Statistical Analysis
The data obtained were analyzed using descriptive statistics parameters, analysis of variance, comparison of means, analysis of correlation and analysis of agglomerative hierarchical clustering (AHC), and the means comparison Student–Newman–Keuls test (SNK) using the statistical analysis software XLSTAT (version 2016).
3. Results
3.1. Analysis of Variability
The analysis of the morphometric parameters studied at the level of the chestnut leaves of each of the seven watersheds (Table 1) allowed us to deduce that the watershed of Azla holds the leaves having the longest petioles (PL,) with an average of 2.43 cm, and the highest LL/LW ratio (2.88). This ratio provides information on the shape of the leaf, and the fact that it is high means that the leaves have a longer shape compared to those from other watersheds. However, the leaves of this watershed are the narrowest (LW) (6.31 cm). They also have the smallest distance between the base and the greatest width (DBW) (8.26 cm), and, finally, they have the smallest blade area (S) (74.35 cm2). The highest coefficient of variation of all the parameters in the seven watersheds was observed for the blade area parameter (S) (34.35%) in the Azla watershed.
The Oued Lakhmiss watershed’s ecotype’s leaves have the shortest blade (LL) (16.84 cm) and the smallest perimeter (38.73 cm). However, Oued Laou watershed’s ecotype’s leaves have the highest perimeter (P) (41.65 cm) and also the longest blade (LL) (18.31 cm). The widest leaves (LW) are those of Tamrabet watershed’s ecotype, with an average width of 7.48 cm. They also have the highest distance between the base and the greatest width (DBW) (9.10 cm), the largest surface (91.42 cm²), the smallest petiole length (PL) (180 cm), and the highest LL/DBW ratio (2.22). It is at the level of this same watershed that the LL/LW ratio varies the least because it has the smallest coefficient of variation for all the parameters at the level of the seven watersheds (6.33%). In the Aouchtem watershed, the leaves have the smallest LL/LW ratio, with an average of 2.32.
The analysis of variance (ANOVA) for the studied morphometric parameters at the leaf level (Table 2) showed a very highly significant variation (p < 0.0001) for all the studied parameters. These results reflect the morphological diversity between the individuals that compose the seven watersheds and indicate that the studied parameters relating to the leaves size and shape of the different ecotypes may be valid indicators for detecting phenotypic variability [2,6,14].
The comparison of the means of the leaves morphometric parameters by the Student–Newman–Keuls (SNK) test at the 5% threshold (Table 3) revealed five modes of grouping between the seven watersheds. Thus, for the perimeter (P) and the lamina length (LL), we did not find any similarity between the means. The petiole length (PL) and the ratio (LL/DBW) allowed us to classify the watersheds into six groups, suggesting the similarity between Tamernount and Aouchtem for PL and between Smir and Oued Lakhmiss for the LL/DBW ratio. As for the blade area (S) and the lamina width (LW), the ecotypes were distributed between five groups by associating the ecotypes of Oued Laou, Tamernount, and Aouchtem in a single group. The LL/LW ratio and the distance from base to greatest width (DBW) grouped the watersheds into four and two groups, respectively.
3.2. Correlation Analysis
The analysis of the correlations between the studied parameters showed that there are links between them, which explains the very strong correlations (Table 4). Indeed, we have noticed the presence of strong correlations between the blade area (S) and the lamina width (LW) (r = 0.924), as well as between the blade area (S) and the distance between the base and the greatest width (DBW) (r = 0.835), and, of less importance, between the blade area (S) and the perimeter (P) (r = 0.754). We also revealed strong correlations between the perimeter (P) and the lamina length (LL) (r = 0.896), as well as between the perimeter (P) and the distance between the base and the greatest width (DBW) (r = 0.610). The blade area (S) is negatively related to the LL/LW ratio (r = −0.505), which provides information on the lamina shape. This means that when the area increases, the ratio decreases, which indicates that the sheet becomes wider than it is long. It was also noted that the petiole length (PL) is negatively correlated with the blade area (S).
3.3. Agglomerative Hierarchical Clustering Analysis (AHC)
Agglomerative hierarchical clustering (AHC) analysis was used to produce a diagram (Figure 3) relating to the analysis of the studied parameters of the leaves of chestnut ecotypes from the seven watersheds. The topology of the dendrogram clearly shows the existence of three large groups relating to the leaves’ studied parameters. The most similar watersheds are Tamernout and Smir on one side and Oued Laou and Aouchtem on the other one.
4. Discussion
The study used the morphometric parameters of 6200 leaves to analyze the variability of Moroccan chestnut ecotypes. The results obtained confirm that the leaves’ morphological traits are very variable. The highest coefficient of variation was observed for the blade area (S) in the Azla watershed, which consists of coppice and natural ecotypes. This same observation was made for Croatian ecotypes [15]. In contrast, the least variable parameter was the shape ratio LL/LW, which is explained by the stability of the shape ratios [6].
Comparing our results with other studies, we notice that leaves from the Azla ecotypes have a blade area (S) (74.35 cm²) almost similar to that of Croatian ecotypes (74.30 cm²) [15]. The petiole length (PL) (2.43 cm), lamina length (LL) (17.73 cm), and lamina width (LW) (6.31 cm) of the Azla ecotypes are very close to the values noted in Portuguese ecotypes [16] of 2.40 cm for petiole length, 17.7 cm for lamina length, and, finally, 6.40 cm for lamina width. The LL/LW ratio (2.88) at this watershed is identical to the one recorded in Bosnia [17].
The leaves of Tamernout ecotypes have a petiole length (PL) (2.28 cm), close to the ecotypes in Portugal (2.20 cm) [16]. Their lamina length (LL) (17.65 cm) is close to Turkish ecotypes (17.63 cm) [3].
The leaves of Oued Lakhmiss ecotypes have a petiole length (PL) (2.18 cm) comparable to Turkish ecotypes (2.11cm) [3]. Their lamina length (LL) (16.84 cm) is close to that of Ukrainian ecotypes (16.7 cm) [18]. The LL/LW ratio (2.45) at this watershed is similar to the ecotype of Bosnia [17].
The leaves of Tamrabet have a petiole length (PL) (1.80 cm) and lamina width (LW) (7.48 cm) almost similar to the values reported in Turkish ecotypes [3], which are around 1.79 cm for the petiole length and 7.40 cm for the lamina width. Their lamina length (LL) (18.11 cm) is close to the ecotype of Portugal (18.10 cm) [17]. At this watershed, the LL/LW ratio (2.61) is also identical to the ecotype of Bosnia [17].
The petiole length (PL) of Oued Laou leaves (2.04 cm) is equal to the ecotype of Bosnia [18]. Lamina length (LL) (18.31 cm) is almost similar to the leaves from Croatian ecotypes (18.30 cm) [15].
The agglomerative hierarchical clustering (AHC) analysis showed the distribution of the seven watersheds in three distinct groups: Tamernout and Smir watersheds on one side and Oued Laou and Aouchtem on the other one, being the closest. The leaves of the Oued Laou and Aouchtem ecotypes are essentially characterized by large values in terms of blade area (S), perimeter (P), lamina length (LL), lamina width (LW), and larger distances between base and width (DBW). They also have small petioles (PL). In comparison, the leaves from the ecotypes of Smir and Tamernout, are distinguished by having small values of perimeter (P), lamina length (LL), and LL/LW and LL/DBW ratios.
Furthermore, the results of the comparison of the means by the Student–Newman–Keuls test (SNK) show a great variability within and between ecotypes that compose the seven watersheds.
5. Conclusions
This study is the first contribution to the knowledge of the ecotypes of the Moroccan chestnut tree. The results obtained show a high degree of variability between and within ecotypes. Thus, the wide range of variability in the morphometric parameters of the leaves confirms that the region is an important center of diversity of chestnut genetic resources. This work was accomplished by the fruit phenotypic characterization. Thus, the evaluation of this diversity is crucial for the implementation of strategies for the conservation and enhancement of this heritage.
Author Contributions
Conceptualization, Y.H., I.T. and S.E.F.; methodology, Y.H., I.T., S.E.F. and M.A.; software, I.T. and S.E.F.; validation, Y.H. and M.A.; formal analysis, I.T. and S.E.F.; investigation, I.T. and Y.H.; resources, Y.H., I.T., S.E.F. and M.A.; data curation, I.T. and Y.H.; writing—original draft preparation, I.T.; writing—review and editing, Y.H. and S.E.F.; visualization, Y.H., I.T., M.A. and S.E.F.; project administration, Y.H., I.T. and S.E.F. All authors have read and agreed to the published version of the manuscript.
Funding
This work was supported by the United Nations Educational Scientific and Cultural Organization [4500409557–506606], and the University Abdelmalek Essâadi scholarship [2624530238].
Data Availability Statement
Not applicable.
Acknowledgments
The authors thank the farmers of the various villages who contributed to the realization of this study, with special thanks to Derdabi M. for his support in the field.
Conflicts of Interest
The authors declare no conflict of interest.
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Figure 1.
Distribution of villages in the seven watersheds at the study area level.
Figure 2.
Diagram representing the measurements of the chestnut tree leaves.
Figure 3.
Grouping dendrogram of the studied ecotypes based on leaf parameters.
Table 1.
The studied morphometric parameters for the leaf in the seven watersheds.
| Watersheds | Parameters | S (cm2) | P (cm) | PL (cm) | LL (cm) | LW (cm) | DBW (cm) | LL/LW | LL/DBW |
|---|---|---|---|---|---|---|---|---|---|
| Oued Laou | Mean | 88.76 | 41.65 | 2.04 | 18.31 | 7.16 | 8.82 | 2.61 | 2.16 |
| SD | 24.81 | 5.53 | 0.38 | 2.56 | 1.26 | 1.51 | 0.40 | 0.21 | |
| CV (%) | 28.54 | 13.28 | 19.77 | 13.99 | 18.29 | 16.15 | 15.33 | 9.51 | |
| Tamernout | Mean | 84.54 | 40.06 | 2.28 | 17.65 | 6.97 | 8.97 | 2.49 | 2.14 |
| SD | 29.70 | 6.79 | 0.41 | 3.35 | 1.33 | 2.25 | 0.38 | 1.17 | |
| CV (%) | 25.58 | 13.98 | 10.23 | 14.72 | 12.88 | 16.5 | 33.08 | 11.08 | |
| Tamrabet | Mean | 91.42 | 41.56 | 1.8 | 18.11 | 7.48 | 9.1 | 2.61 | 2.22 |
| SD | 32.22 | 7.20 | 0.73 | 3.42 | 1.58 | 2.09 | 0.54 | 0.64 | |
| CV (%) | 15.45 | 7.68 | 14.24 | 8.73 | 9.05 | 13.17 | 6.33 | 28.92 | |
| Aouchtem | Mean | 88.95 | 40.34 | 2.05 | 17.35 | 7.45 | 8.85 | 2.32 | 1.98 |
| SD | 32.22 | 8.29 | 0.40 | 4.27 | 1.36 | 2.42 | 0.42 | 0.20 | |
| CV (%) | 22.06 | 12.65 | 8.97 | 13.22 | 9.39 | 16.2 | 17.87 | 10.27 | |
| Azla | Mean | 74.35 | 39.68 | 2.43 | 17.73 | 6.31 | 8.26 | 2.88 | 2.17 |
| SD | 30.99 | 8.66 | 0.65 | 4.17 | 1.39 | 2.11 | 0.68 | 0.19 | |
| CV (%) | 34.35 | 19.03 | 20.57 | 20.26 | 16.73 | 22.07 | 23.58 | 8.84 | |
| Oued Lakhmiss | Mean | 79.82 | 38.73 | 2.18 | 16.84 | 7 | 8.28 | 2.45 | 2.10 |
| SD | 20.12 | 5.20 | 0.37 | 2.50 | 0.96 | 1.48 | 0.32 | 0.20 | |
| CV (%) | 25.43 | 13.24 | 18.03 | 14.7 | 14.4 | 17.64 | 13.03 | 9.33 | |
| Smir | Mean | 84.34 | 39.64 | 2.21 | 17.53 | 7.20 | 9.03 | 2.47 | 1.97 |
| SD | 21.23 | 4.83 | 0.33 | 2.24 | 1.09 | 1.30 | 0.26 | 0.11 | |
| CV (%) | 25.17 | 12.18 | 14.82 | 12.76 | 15.21 | 14.41 | 10.73 | 5.71 |
SD: standard deviation CV: coefficient of variation.
Table 2.
Analysis of variance (ANOVA) of the studied morphometric parameters of the leaves in the ecotypes of Castanea sativa Mill.
Table 2.
Analysis of variance (ANOVA) of the studied morphometric parameters of the leaves in the ecotypes of Castanea sativa Mill.
| Parameters | S | P | PL | LL | LW | DBW | LL/LW | LL/DBW |
|---|---|---|---|---|---|---|---|---|
| F | 31.55 | 29.72 | 16.799 | 29.49 | 37.32 | 32.62 | 28.91 | 16.34 |
| Pr > F | <0.0001 **** | <0.0001 **** | <0.0001 **** | <0.0001 **** | <0.0001 **** | <0.0001 **** | <0.0001 **** | <0.0001 **** |
**** very highly significant test.
Table 3.
Comparison of the means of the leaves’ studied morphometric parameters in chestnut ecotypes by Student–Newman–Keuls test (SNK).
Table 3.
Comparison of the means of the leaves’ studied morphometric parameters in chestnut ecotypes by Student–Newman–Keuls test (SNK).
| Watersheds | S | P | PL | LL | LW | DBW | LL/LW | LL/DBW |
|---|---|---|---|---|---|---|---|---|
| Tamrabet | A | A | A | A | A | A | A | A |
| Oued Laou | B | B | B | B | B | A | B | B |
| Tamernount | B | C | C | C | B | A | B | C |
| Aouchtem | B | D | C | D | B | A | C | D |
| Azla | C | E | D | E | C | A | C | E |
| Smir | D | F | E | F | D | B | C | F |
| Oued Lakhmiss | E | G | F | G | E | B | D | F |
Table 4.
Matrix of the correlation coefficients of the leaves’ studied parameters.
| Variables | P | PL | LL | LW | DBW | LL/LW | LL/DBW |
|---|---|---|---|---|---|---|---|
| S | 0.754 | −0.877 | 0.446 | 0.924 | 0.835 | −0.505 | −0.025 |
| P | 1 | −0.681 | 0.896 | 0.472 | 0.610 | 0.154 | 0.438 |
| PL | 1 | −0.350 | −0.867 | −0.579 | 0.337 | −0.169 | |
| LL | 1 | 0.104 | 0.480 | 0.491 | 0.546 | ||
| LW | 1 | 0.741 | −0.723 | −0.272 | |||
| DBW | 1 | −0.399 | −0.119 | ||||
| LL/LW | 1 | 0.677 |
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Toujgani, I.; El Fatehi, S.; Ater, M.; Hmimsa, Y. Phenotypic Variation of Castanea sativa Mill. Ecotypes in Northern Morocco Based on a Multivariate Analysis of Leaf Morphometrics. Biol. Life Sci. Forum 2021, 2, 34. https://doi.org/10.3390/BDEE2021-09458
AMA Style
Toujgani I, El Fatehi S, Ater M, Hmimsa Y. Phenotypic Variation of Castanea sativa Mill. Ecotypes in Northern Morocco Based on a Multivariate Analysis of Leaf Morphometrics. Biology and Life Sciences Forum. 2021; 2(1):34. https://doi.org/10.3390/BDEE2021-09458
Chicago/Turabian StyleToujgani, Ihssane, Salama El Fatehi, Mohammed Ater, and Younes Hmimsa. 2021. "Phenotypic Variation of Castanea sativa Mill. Ecotypes in Northern Morocco Based on a Multivariate Analysis of Leaf Morphometrics" Biology and Life Sciences Forum 2, no. 1: 34. https://doi.org/10.3390/BDEE2021-09458
APA StyleToujgani, I., El Fatehi, S., Ater, M., & Hmimsa, Y. (2021). Phenotypic Variation of Castanea sativa Mill. Ecotypes in Northern Morocco Based on a Multivariate Analysis of Leaf Morphometrics. Biology and Life Sciences Forum, 2(1), 34. https://doi.org/10.3390/BDEE2021-09458
