Accessing agro-morphological variation in Ber (Ziziphus spp.) Germplasm from Kutch, Gujarat, Using Multivariate Techniques

Accessing agro-morphological variation in Ber (Ziziphus spp.) Germplasm from Kutch, Gujarat, Using Multivariate Techniques | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Accessing agro-morphological variation in Ber (Ziziphus spp.) Germplasm from Kutch, Gujarat, Using Multivariate Techniques Rahul Dev, Traloki Singh, Chandan Kumar, Arvind Tetarwal, Devi Dayal This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8618673/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 17 You are reading this latest preprint version Abstract This study assessed morphological diversity in ber ( Ziziphus spp. ) germplasm collected from Gujarat’s Kutch district. Thirty accessions from Bhuj, Nakhatrana, Mandvi, and Abdasa talukas were sampled across ~ 170 km, capturing the central and western Kutch ber-growingarea. Seventeen morphological and quality traits including plant height, canopy spread, branch number, leaf area, fruit dimensions, weight and total soluble solids (TSS) were measured. Descriptive statistics showed broad phenotypic dispersion: plant height ranged from 144 to 455 cm; canopy volume varied from 0.58–8.34 m³; and leaf area from 1.38–9.76 cm², TSS ranged from 3.40–17.50 °Brix. Fruit length, width, and 5-fruit weight displayed moderate variability. Principal component analysis reduced the 17 traits to four components, explaining 85.55% of total variation. PC1 (50.14%) was associated with fruit dimensions and TSS, identifying high-yield, high-quality accessions (e.g., Ber_Abda_20, Ber_Bhuj_30); PC2 (20.50%) contrasted vegetative vigour with fruit size. Hierarchical clustering separated accessions into four clusters: (1) tall, large-fruited types; (2) a unique accession combining moderate vigour and very large, sweet fruits; (3) compact, small-fruited types; and (4) intermediate vigour and fruit size. Correlation analysis revealed strong positive associations between plant height and canopy volume (r ≈ 0.91) and between fruit weight and size index (r ≈ 0.58), while TSS correlated moderately with plant size and fruit dimensions. Overall, the Kutch ber germplasm displayed considerable morphological and biochemical diversity. The promising accessions Ber_Abda_20, Ber_Bhuj_21, Ber_Bhuj_29 and Ber_Bhuj_30 offer potential for cultivar development and highlight the value of conserving Kutch’s under-studied genetic resources. Ziziphus mauritiana germplasm diversity Kutch district morphological traits principal component analysis cluster analysis fruit quality. Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Introduction India's jujube, or ber ( Ziziphusmauritiana Lam ) has been grown for thousands of years throughout the Indo-Pak subcontinent, as well as much of Asia and Africa, as an evergreen fruit tree belonging to the Rhamnaceae family. There are between 50 and 170 species in the genus Ziziphus , but only a handful especially Z . Mauritiana , Z . jujuba as well as Z . For their edible drupes, nummularia are widely cultivated (Anjum et al. , in 2018). While Z. mauritiana is widely grown in India, Z . jujuba is the most common in China, and Z. nummularia grows as a shrub in arid areas (Sareen et al. , 2023). Because of its deep taproot and capacity to go dormant during drought, the tree can withstand high temperatures and saline or alkaline soils and flourishes in hot, dry climates (Kumar and Tripathi, 2024 ). These characteristics make ber especially suited to the arid and semiarid regions of Rajasthan, Gujarat, Uttar Pradesh and other Indian states (Kumar et al., 2021 ). Ber trees show considerable morphological variation. Wild trees may be 1.5–2 m tall shrubs, while improved cultivars can reach 10–15 m with spreading crowns and thorny branches. Leaf shape ranges from ovate to elliptic, and flowers are small, yellow, and borne in axillary clusters (Kumar and Tripathi, 2024 ). Ber trees vary greatly in their morphology. The height of wild trees can range from 1.5 to 2 meters, whereas improved cultivars with spreading crowns and robust branches can grow up to 10 to 15 meters. Leaf shape ranges from ovate to elliptic, and flowers are small, yellow, and borne in axillary clusters (Kumar and Tripathi, 2024 ). The fruit can be round, oblong or oval in size, ranging from 2 to 3 cm in wild forms to more than 5 cm in the cultivated species (Anjum et al., 2018 ) After ripening, the colour changes from green to yellow or red brown and the meat may be crisp or mealy. Indian ber trees are cross pollinated; this promotes wide genetic variation, as evidenced by the many landraces in India and Pakistan (Kumar and Tripathi, 2024 ; Kumar et al., 2021 ). Wild Z . nummularia is distinctly different from cultivated ber in that it produces small, spiny shrubs with small leaves and fruits, but provides fodder and rootstock material to local economies (Sareen et al. , 2023). Nutritionally, ber fruits are rich in carbohydrates, sugars, dietary fibre, vitamin C, vitamin A, and minerals such as calcium and phosphorus (Anjum et al., 2018 ). They contain phenolics, flavonoids and other antioxidants (Anjum et al., 2018 ); wild jujube ( Z. nummularia ) has been noted for its high vitamin C content and medicinal uses in treating skin infections, bronchitis and digestive disorders (Sharif et al., 2022 ). Leaves are valued as fodder for livestock, and the wood serves as fuel and for making implements (Anjum et al., 2018 ). Because of their nutritional density and resilience under marginal conditions, ber fruits are sometimes called the “poor man’s apple”. Ber fruit is nutritionally rich in carbohydrates, sugars, dietary fibres, vitamins C and A and minerals such as calcium and phosphorus (Anjum et al., 2018 ). They contain phenols, flavonoids and other antioxidants and have been reported to be used in the treatment of skin infections, bronchitis and digestive disorders (Sharif et al., 2022 ). The leaves are valued as feed for cattle and the wood is used as fuel and for the manufacture of tools. Due to its high nutritional value and its resilience to harsh conditions, ber fruits are sometimes referred as the poor man's apple (Anjum et al., 2018 ). Despite its economic and nutritional value, the use of ber is still under-explored. Nutrition analysis of the exogenous samples from Ziziphus species in the arid areas of Rajasthan revealed a high variability of phenols, antioxidants and ascorbic acid in the genotypes (Meena, 2023).Studies in Bundelkhand and East Uttar Pradesh have shown large differences in the weight (6–30 g), length (1.6–4.4 cm), total soluble solids (14–20 brix) and ascorbic acid among ber accessions (Kumar et al., 2021 ). Evaluation of indigenous ber germplasm in Jammu region has shown significant differences in the ratio of pulp to stone, juice and sugar profiles (Kumari, 2015). However, wild diversity in the ber is still poorly characterised and is threatened by habitat loss, overgrazing and the spread of improved cultivars (Sharif et al., 2022 ). Information on seed characteristics such as seed weight and germination is also limited for the rootstock species on which the berry is grown (Srivastava et al., 2001 ). Germplasm from Punjab, Rajasthan, and Uttar Pradesh, ex-situ collections have been studied, but the dry Kachchh (Kutch) district of Gujarat, which is severely droughted and salinity has been neglected. Because ber in Kutch has evolved under extreme conditions, its landraces may have special characteristics for drought tolerance, salinity resistance, and fruit quality. Systematic characterization of this germplasm (morphological, agronomic and physiochemical traits) is needed to identify superior genotypes and to preserve valuable diversity for future breeding and cultivation. Material and methods Field survey and germplasm collection During the summer (February–March) season of 2019, field surveys were conducted in five talukas in arid Kachchh, Gujarat: Bhuj, Mandvi, Nakhatarana, Bhachau and Rapar. The purpose of these surveys was to collect Ziziphus mauritiana germplasm from various habitats (Table 1). In carrying out field surveys, passport information on the collection site, the associated vegetation and the conservation status of the species has been recorded. Passport information was also collected during the germplasm collection and is shown in Fig. 1. During the exploration visits, information on various growth parameters, including plant height (cm), plant spread (N-S), plant spread (E-W), number of branches/plant, and stem girth (mm), were collected in the field for morphological characterisation. Following the tour, twenty fruits from each genotype were chosen at random, and measurements of the fruits' thickness (mm), length (mm), width (mm), No. of fruit per 100 g and TSS (º) were made in the lab. Twenty fruit samples were measured for length, width and thickness using a digital caliper with a precision of 0.01 mm. Additional morphological characteristics include Petiole length (mm), Petiole thickness (mm), Leaf area (cm2), and based on the leaf that was collected. The size, shape and volume of the fruit were described by the main morphometric parameters calculated from these measurements. A simple indicator of total fruit size, the Fruit Size Index (FSI) is computed as the product of fruit width and length (FSI = Length × Width). The Fruit Shape Index measures how elongated a fruit is by dividing its length by its width (Shape Index = Length / Width). A round shape is suggested by values near 1, whereas elongated or flattened forms are indicated by values much greater or less than 1. The fruit's degree of symmetry is expressed by the Fruit Roundness or Elongation Ratio (Roundness = Smaller dimension / Larger dimension × 100), where higher percentages indicate more spherical fruits. Fruit Volume (approximate), the following formula was used to estimate fruit volume, assuming an ellipsoidal geometry. Where L is fruit length and W is fruit width. The surface area (approximate) was estimated using the ellipsoidal formula developed by Knud Thomsen. Where a = L/2, b = c = W/2, and p ≈ 1.6. Canopy volume for each tree was estimated with the “contour method” (Wright et al. , 2005), treating the canopy as a geometric solid whose base widths ‘a’ (perpendicular to the planting row) and ‘b’ (parallel to the row) were measured at the canopy base, and whose height ‘h’ was taken from the lowest branch to the apex. The contour functions ‘m(x)’ and ‘m(y)’ adjust for departures from an ideal canopy shape. The final expression is (CV = [(1⁄4) π a b h) / (m(x) + m(y) + 1]. Data Analysis All morphological characteristics of 30 wild Ber ( Ziziphus spp .) accessions have been analysed for multivariate relationships between accessions. Principal component analysis (PCA) was performed to identify the traits that contribute the most to inter-genotypic variation, and a biplot of the first two PCs was used to visualise genotypic dispersion and trait loading. Hierarchical clustering has been calculated using Ward's minimum variance algorithm (Ward's D²) for Euclidean distances. Pearson's correlation coefficients were used to quantify pairwise trait associations, and a heatmap was created to illustrate both positive and negative relationships that are pertinent to selection. Python (v3.11) was used for all analyses, with Matplotlib/Seaborn for visualisation, SciPy for hierarchical clustering, and scikit-learn for PCA and scaling. Results and Discussion Collection and survey results In February and March 2019, systematic field surveys were conducted in the Kutch district of Gujarat to collect 30 geo-referencedber ( Ziziphus spp.) accessions from the talukas of Bhuj, Nakhatrana, Mandvi, and Abdasa. The distribution of accessions was as follows: Mandvi (n = 4; 13.33%), Bhuj (n = 17; 56.67%), Nakhatrana (n = 8; 26.67%), and Abdasa (n = 1; 3.33%), with multiple sampling sites covered within each taluka (Table 1). The surveyed area spanned approximately 170 km, with the centroid located at 23.3688° N, 69.5494° E, extending from 23.0347–23.6881° N and 69.2051–70.7156° E. Three accessions represented the geographical extremes: Br_Mand_7 (southernmost; 23.0347° N, 69.3000° E), Br_Nakh_29 (northernmost and westernmost; 23.6881° N, 69.2051° E), and Br_Mand_8 (easternmost; 23.5629° N, 70.7156° E) (Fig. 1). Together, this coverage encompassed the major ber-growing tracts of central and western Kutch, ensuring adequate representation both within and across talukas for subsequent diversity analyses.Comparable botanical surveys in the Indian arid zone have similarly emphasized broad geographic coverage to capture genetic variability. For instance, (Kumar and Tripathi, 2024 ) documented 40 ber genotypes from eastern Uttar Pradesh (Dev et al., 2023 ) reported 27 genotypes of Grewia tenax and 16 genotypes of Cordia gharaf from the Kachchh region of Gujarat; while (Venkatesan et al ., 2018) collected 40 Grewia tenax genotypes from Barmer, Jaisalmer, Jodhpur, and Pali districts of western Rajasthan. Descriptive Statistics A wide range of phenotypic variation was observed in both fruit and plant architectural traits. Plant height varied from 144 to 455 cm, with an average of 255.25 ± 77.49 cm (CV = 30.36%). The tallest plant was Ber_Bhuj_30, while the shortest was Ber_Bhuj_1. Similar variability in plant height has been reported in other germplasm surveys; for example, Kumar and Tripathi ( 2024 ) documented heights ranging from 3.22 to 5.50 m across 40 genotypes, while Kumar et al., ( 2021 ) reported heights between 1.6 and 4.4 m. Significant differences were also recorded in plant spread. The north south spread ranged from 132 to 525 cm (CV = 32.13%), whereas the east–west spread varied between 147 and 540 cm (CV = 36.66%), with Ber_Abda_20 showing the widest east–west spread. Canopy volume exhibited the greatest variability, ranging from 0.58 to 8.34 m³ (CV = 66.24%). The smallest canopy volume was observed in Ber_Bhuj_1 and the largest in Ber_Bhuj_30. Such wide variation in canopy traits has also been highlighted in other ber germplasm evaluations Kumar and Tripathi, ( 2024 ). Branching habits showed substantial variation, with the number of primary branches per plant ranging from 2 to 20 (CV = 62.19%). Leaf area also varied widely, from 1.38 to 9.76 cm². Leaf size is a key parameter influencing photosynthesis, as well as food, fodder, and biomass production. Comparable variation has been reported in Grewia tenax germplasm from Kutch, Gujarat, where leaf size ranged from 3.1 to 8.3 cm² (mean 4.8 cm²; Dev et al., ( 2023 ). Similar variability in leaf shape (cordate, oval, obovate and elliptic) and other morphological traits has also been documented in ber genotypes from eastern Uttar Pradesh (Kumar and Tripathi, 2024 ). Such phenotypic diversity in morpho-physico-chemical traits has likewise been observed in other crops, as reported by Özrenk et al., ( 2010 ), Karadeniz et al., ( 2013 ); Dev et al., ( 2025 ). This diversity is likely influenced by multiple factors, including geographical distribution, origin, genotype, climatic conditions, and their complex interactions. Fruit Traits For the selection of superior genotypes, fruit characteristics particularly fruit weight are among the most important target traits in ber improvement programs. In the present study, fruit size attributes varied from small to large types. The mean fruit length was 10.67 mm (range: 8.29–13.05 mm; CV = 10.14%) and the mean width was 11.27 mm (range: 8.77–14.73 mm; CV = 10.69%). The composite fruit size (length × width) was smallest in Ber_Bhuj_2 and largest in Ber_Abda_20 (Table 2). Similar variability in fruit weight, 100‑seed weight, fruit length, and width has been documented in ber and other horticultural crops. For instance, such variation has been reported in ber (Kumar et al., 2021 ; Sharif et al., 2022 ; Meena et al., 2023 ), Grewiatenax (Dev et al., 2023 ), apple (Miller et al., 2004 ; Kaya et al., 2015 ), fig (Darjazi, 2011), apricot (Kumar, 2015), and Cordia gharaf (Dev et al., 2024 ). In the present collection, fruit roundness, surface area, and fruit shape index were the most consistent attributes across genotypes. Fruit test weight (308–1496 g), five‑fruit weight (1.54–7.48 g), and fruit volume (353.20–1332.10 mm³) showed comparable variability (CV ≈ 29–32%), with the highest values consistently observed in Ber_Abda_20. Similar significant differences in fruit size and weight have also been reported in indigenous jujube cultivars of Pakistan, where the cultivar Foladi exhibited comparatively high fruit weight and width (Anjum et al., 2018 ). Previous diversity studies have emphasized that variation in morphological traits provides a reliable basis for germplasm selection and improvement (Meghwal et al., 2014 ; Kumar et al., 2015 ; Dev et al., 2025 ). Total soluble solids (TSS) displayed high variability (CV = 37.99%), ranging from 3.40 to 17.50 °Brix, with Ber_Bhuj_21 recording the highest value. Similar variation (5.9–13.2 °Brix) was also noted among wild ber genotypes in Pakistan (Sharif et al., 2022 ). Among Indian cultivars, TSS values ranged more widely (12.5–24.2 °Brix), which may be attributed to genetic differences, cultural practices, and stage of ripening Anjum et al., ( 2018 ). Consistent with this trend, cultivars such as Khobani (14.92 °Brix) and Dilbahar (11.33 °Brix) showed contrasting TSS levels, Anjum et al., ( 2018 ); Kumari et al. ( 2015 ) further reported that the cultivar Sanaur‑4 recorded high TSS (15.76 °Brix) and a superior TSS/acid ratio (40.38), along with distinct sugar profiles and vitamin C content across genotypes. In the current study, canopy volume, branching pattern, and stem girth were among the most variable traits, whereas fruit roundness and shape remained relatively stable across accessions. Based on a combination of vigour, fruit size, and quality attributes, the accessions Ber_Abda_20, Ber_Bhuj_21, Ber_Bhuj_29, and Ber_Bhuj_30 were identified as the most promising candidates for further evaluation and potential use in crop improvement programs. Principal Component Analysis Principal component analysis (PCA) reduced the original 17 traits to four principal components (PCs) with eigenvalues greater than 1, together explaining 85.55% of the total variation. PC1 (eigenvalue = 8.817; 50.14% variance) showed strong positive loadings for fruit width, fruit length, five‑fruit weight, size index, fruit volume, fruit surface area, test weight, and total soluble solids (TSS). This axis represented accessions combining large fruit size with improved quality traits, such as Ber_Abda_20 and Ber_Bhuj_30. PC2 (eigenvalue = 3.605; 20.50% variance) contrasted vegetative vigour against fruit traits (Fig. 2). High positive loadings were recorded for plant height, canopy spread, stem girth, canopy volume, and branch number, whereas fruit length, width, size index, shape index, and roundness loaded negatively. This axis separated tall, vigorous types (e.g., Ber_Bhuj_30, Ber_Bhuj_21) from compact, fruit-dominant accessions. PC3 (eigenvalue = 1.515; 8.62% variance) was associated primarily with fruit shape traits (shape index and roundness), while PC4 (eigenvalue = 1.107; 6.29%) highlighted leaf area and stem girth (Fig. 3). Similar PCA findings in ber have shown that the first two PCs often capture a major share of variation, typically separating yield‑related traits from fruit quality attributes Anjum et al., ( 2018 ); Kumar et al., ( 2015 ) reported that the first two PCs explained 62.97% of total variation with one group characterized by superior yield traits and the other by fruit quality. Comparable patterns have also been described in rose germplasm, where fruit diameter, fruit weight, pulp weight, and seed number explained substantial proportions of the total variation (Verma, 2015). Hierarchical Cluster Analysis Using a Euclidean distance dendrogram, four distinct clusters were identified. Cluster 1 consisted of high-vigor, large-fruited types (Ber_Bhuj_21, Ber_Nakh_22, Ber_Bhuj_24, Ber_Bhuj_29, and Ber_Bhuj_30) characterized by tall plants, broad canopies, and large fruits. Cluster 2 included a single accession (Ber_Abda_20) that exhibited moderate vigor combined with extremely large fruits and high total soluble solids (TSS). Cluster 3 comprised eight compact accessions with low vigor and small, round fruits (e.g., Ber_Bhuj_1, Ber_Bhuj_2). Cluster 4 contained 16 accessions with intermediate vigour and fruit size (Fig. 4). Similar grouping patterns based on pomological traits have been reported in cluster analyses of indigenous Indian jujube Anjum et al., ( 2018 ), Wild Ber Sharif et al., ( 2022 ), domesticated and wild Ber Sharif et al., ( 2019 ), and Wild Indian Ber Sareen et al. , (2023). Correlation Analysis The correlation analysis revealed that plant height was strongly correlated with canopy volume (r ≈ 0.91), east–west spread (r ≈ 0.66), and north–south spread (r ≈ 0.73). Fruit weight (measured from 5 fruits) showed positive correlations with fruit width (r ≈ 0.47), size index (r ≈ 0.58), volume (r ≈ 0.61), surface area (r ≈ 0.61), and test weight (r ≈ 0.82). The shape index (length/width) was negatively correlated with width (r ≈ − 0.40), size index (r ≈ − 0.46), and surface area (r ≈ − 0.40). Total soluble solids (TSS) demonstrated moderate positive correlations with plant height, canopy spreads, and fruit dimensions (Fig. 5). Similar correlations have been reported in wild jujube populations, where leaf length and width showed a strong positive correlation (r = 0.897), and fruit weight was negatively correlated with stone width (r = − 0.409) Sharif et al., ( 2022 ), supporting the present findings. Significant positive correlations were also observed for leaf dimensions, fruit weight, and fruit size (Azam-Ali et al., 2001 ). Conclusion The study surveyed and collected 30 geo-referenced ber accessions from the Kutch district, revealing significant phenotypic and genetic diversity in plant and fruit traits. Multivariate analyses identified four distinct clusters, highlighting promising accessions like Ber_Abda_20 and Ber_Bhuj_30 for breeding. Strong correlations between vegetative growth and fruit quality traits were observed, providing valuable insight for future ber improvement and conservation in arid regions of Gujarat. Declarations Funding Declaration This research did not receive any specific grant from funding agencies in the public, commercial, or not-for-profit sectors. All costs associated with the research and manuscript preparation were covered by the authors' institution. Permissions to collect the plants/plant parts The plant materials used in this study were collected from non-protected areas and involved non-endangered, commonly cultivated species. The species investigated is a commonly cultivated crop, and no specific permits or licenses were required for collection. Data Availability Statement The datasets generated during and/or analysed during the current study are available from the corresponding author on reasonable request. Ethics and Consent to Participate The plant material used in this study was obtained from cultivated sources. All procedures involving plant collection complied with relevant local and national guidelines and regulations. Samples were collected from non-protected areas, and all relevant institutional and regulatory requirements were followed. Author Contribution R.D. write the manuscriptT.S., C.K., A.T., D.D. collection and analyses of Data, All authors reviewed the manuscript. 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Additional Declarations No competing interests reported. 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Institute","correspondingAuthor":false,"prefix":"","firstName":"Traloki","middleName":"","lastName":"Singh","suffix":""},{"id":601830738,"identity":"53599dbd-577d-40f2-8bb5-65bcd4132cea","order_by":2,"name":"Chandan Kumar","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABAElEQVRIiWNgGAWjYDACCSDmgdIMDBUScAkeIrWcIVkLYxsR7uKf3fzswZuaOwySM7ITP1fOs5Dnn32A8XHFLwYZc1yW3Dlmbjjn2DMGaYnczZJnt0kYzjiXwGx4to+Bx7IBuxYDiQQzaR62wwxyErkbJBu3SSQwnGFgk2zsYeAxOIBLS/o3aZ5/YC2bfzbOkUiQJ6wlx0yat+0wyGHbJBsbJBIMQFoafuDWInEjp0xybt9hHsmet9ssG45JGG48w9hsCNSLUwv/jPRtEm++HZaTOJ67+WZDTZ283Bnmgw8b/tjY49ICA8gRx9gAjCAJnEpxgT8k6xgFo2AUjILhCwAQOFPJH5OFPwAAAABJRU5ErkJggg==","orcid":"","institution":"Central Arid Zone Research Institute","correspondingAuthor":true,"prefix":"","firstName":"Chandan","middleName":"","lastName":"Kumar","suffix":""},{"id":601830739,"identity":"edfa3731-c911-4cdd-9c00-4acebb145e5b","order_by":3,"name":"Arvind Tetarwal","email":"","orcid":"","institution":"Central Arid Zone Research Institute","correspondingAuthor":false,"prefix":"","firstName":"Arvind","middleName":"","lastName":"Tetarwal","suffix":""},{"id":601830740,"identity":"9331d381-da6b-464c-b91b-f54ce28e0299","order_by":4,"name":"Devi Dayal","email":"","orcid":"","institution":"Central Arid Zone Research Institute","correspondingAuthor":false,"prefix":"","firstName":"Devi","middleName":"","lastName":"Dayal","suffix":""}],"badges":[],"createdAt":"2026-01-16 12:05:06","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-8618673/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-8618673/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":104404997,"identity":"20f7a28b-4a19-44a4-a890-1934dc14a609","added_by":"auto","created_at":"2026-03-11 12:21:32","extension":"jpg","order_by":1,"title":"Figure 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21:20:34","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":14128,"visible":true,"origin":"","legend":"","description":"","filename":"Table1Collectionsites.docx","url":"https://assets-eu.researchsquare.com/files/rs-8618673/v1/3f9e99de7b840be025a74e37.docx"},{"id":104405508,"identity":"0cac37c2-2e8e-48e3-9b54-150d1b183a95","added_by":"auto","created_at":"2026-03-11 12:23:09","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":21138,"visible":true,"origin":"","legend":"","description":"","filename":"Table2Growthdata.docx","url":"https://assets-eu.researchsquare.com/files/rs-8618673/v1/08be2b798384da544084d3ea.docx"},{"id":104268690,"identity":"07bc2019-9bec-4f0d-9259-12feda5d7fbf","added_by":"auto","created_at":"2026-03-09 21:20:34","extension":"docx","order_by":3,"title":"","display":"","copyAsset":false,"role":"supplement","size":16909,"visible":true,"origin":"","legend":"","description":"","filename":"Table3PCAloadings.docx","url":"https://assets-eu.researchsquare.com/files/rs-8618673/v1/a6d6921aa6154716e848ad57.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Accessing agro-morphological variation in Ber (Ziziphus spp.) Germplasm from Kutch, Gujarat, Using Multivariate Techniques","fulltext":[{"header":"Introduction","content":"\u003cp\u003eIndia's jujube, or ber (\u003cem\u003eZiziphusmauritiana Lam\u003c/em\u003e) has been grown for thousands of years throughout the Indo-Pak subcontinent, as well as much of Asia and Africa, as an evergreen fruit tree belonging to the Rhamnaceae family. There are between 50 and 170 species in the genus \u003cem\u003eZiziphus\u003c/em\u003e, but only a handful especially \u003cem\u003eZ\u003c/em\u003e. \u003cem\u003eMauritiana\u003c/em\u003e, \u003cem\u003eZ\u003c/em\u003e. \u003cem\u003ejujuba\u003c/em\u003e as well as \u003cem\u003eZ\u003c/em\u003e. For their edible drupes, \u003cem\u003enummularia\u003c/em\u003e are widely cultivated (Anjum \u003cem\u003eet al.\u003c/em\u003e, in 2018). While Z. \u003cem\u003emauritiana\u003c/em\u003e is widely grown in India, \u003cem\u003eZ\u003c/em\u003e. \u003cem\u003ejujuba\u003c/em\u003e is the most common in China, and \u003cem\u003eZ. nummularia\u003c/em\u003e grows as a shrub in arid areas (Sareen \u003cem\u003eet al.\u003c/em\u003e, 2023). Because of its deep taproot and capacity to go dormant during drought, the tree can withstand high temperatures and saline or alkaline soils and flourishes in hot, dry climates (Kumar and Tripathi, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). These characteristics make ber especially suited to the arid and semiarid regions of Rajasthan, Gujarat, Uttar Pradesh and other Indian states (Kumar et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Ber trees show considerable morphological variation. Wild trees may be 1.5\u0026ndash;2 m tall shrubs, while improved cultivars can reach 10\u0026ndash;15 m with spreading crowns and thorny branches. Leaf shape ranges from ovate to elliptic, and flowers are small, yellow, and borne in axillary clusters (Kumar and Tripathi, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Ber trees vary greatly in their morphology. The height of wild trees can range from 1.5 to 2 meters, whereas improved cultivars with spreading crowns and robust branches can grow up to 10 to 15 meters. Leaf shape ranges from ovate to elliptic, and flowers are small, yellow, and borne in axillary clusters (Kumar and Tripathi, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The fruit can be round, oblong or oval in size, ranging from 2 to 3 cm in wild forms to more than 5 cm in the cultivated species (Anjum et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) After ripening, the colour changes from green to yellow or red brown and the meat may be crisp or mealy. Indian ber trees are cross pollinated; this promotes wide genetic variation, as evidenced by the many landraces in India and Pakistan (Kumar and Tripathi, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Kumar et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Wild \u003cem\u003eZ\u003c/em\u003e. \u003cem\u003enummularia\u003c/em\u003e is distinctly different from cultivated ber in that it produces small, spiny shrubs with small leaves and fruits, but provides fodder and rootstock material to local economies (Sareen \u003cem\u003eet al.\u003c/em\u003e, 2023). Nutritionally, ber fruits are rich in carbohydrates, sugars, dietary fibre, vitamin C, vitamin A, and minerals such as calcium and phosphorus (Anjum et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). They contain phenolics, flavonoids and other antioxidants (Anjum et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e); wild jujube (\u003cem\u003eZ. nummularia\u003c/em\u003e) has been noted for its high vitamin C content and medicinal uses in treating skin infections, bronchitis and digestive disorders (Sharif et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Leaves are valued as fodder for livestock, and the wood serves as fuel and for making implements (Anjum et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Because of their nutritional density and resilience under marginal conditions, ber fruits are sometimes called the \u0026ldquo;poor man\u0026rsquo;s apple\u0026rdquo;. Ber fruit is nutritionally rich in carbohydrates, sugars, dietary fibres, vitamins C and A and minerals such as calcium and phosphorus (Anjum et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). They contain phenols, flavonoids and other antioxidants and have been reported to be used in the treatment of skin infections, bronchitis and digestive disorders (Sharif et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The leaves are valued as feed for cattle and the wood is used as fuel and for the manufacture of tools. Due to its high nutritional value and its resilience to harsh conditions, ber fruits are sometimes referred as the poor man's apple (Anjum et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Despite its economic and nutritional value, the use of ber is still under-explored. Nutrition analysis of the exogenous samples from \u003cem\u003eZiziphus\u003c/em\u003e species in the arid areas of Rajasthan revealed a high variability of phenols, antioxidants and ascorbic acid in the genotypes (Meena, 2023).Studies in Bundelkhand and East Uttar Pradesh have shown large differences in the weight (6\u0026ndash;30 g), length (1.6\u0026ndash;4.4 cm), total soluble solids (14\u0026ndash;20 brix) and ascorbic acid among ber accessions (Kumar et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Evaluation of indigenous ber germplasm in Jammu region has shown significant differences in the ratio of pulp to stone, juice and sugar profiles (Kumari, 2015). However, wild diversity in the ber is still poorly characterised and is threatened by habitat loss, overgrazing and the spread of improved cultivars (Sharif et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Information on seed characteristics such as seed weight and germination is also limited for the rootstock species on which the berry is grown (Srivastava et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). Germplasm from Punjab, Rajasthan, and Uttar Pradesh, ex-situ collections have been studied, but the dry Kachchh (Kutch) district of Gujarat, which is severely droughted and salinity has been neglected. Because ber in Kutch has evolved under extreme conditions, its landraces may have special characteristics for drought tolerance, salinity resistance, and fruit quality. Systematic characterization of this germplasm (morphological, agronomic and physiochemical traits) is needed to identify superior genotypes and to preserve valuable diversity for future breeding and cultivation.\u003c/p\u003e"},{"header":"Material and methods","content":"\u003cdiv id=\"Sec3\"\u003e\n \u003ch2\u003eField survey and germplasm collection\u003c/h2\u003e\n \u003cp\u003eDuring the summer (February\u0026ndash;March) season of 2019, field surveys were conducted in five talukas in arid Kachchh, Gujarat: Bhuj, Mandvi, Nakhatarana, Bhachau and Rapar. The purpose of these surveys was to collect \u003cem\u003eZiziphus mauritiana\u003c/em\u003e germplasm from various habitats (Table\u0026nbsp;1). In carrying out field surveys, passport information on the collection site, the associated vegetation and the conservation status of the species has been recorded. Passport information was also collected during the germplasm collection and is shown in Fig.\u0026nbsp;1. During the exploration visits, information on various growth parameters, including plant height (cm), plant spread (N-S), plant spread (E-W), number of branches/plant, and stem girth (mm), were collected in the field for morphological characterisation.\u003c/p\u003e\n \u003cp\u003eFollowing the tour, twenty fruits from each genotype were chosen at random, and measurements of the fruits\u0026apos; thickness (mm), length (mm), width (mm), No. of fruit per 100 g and TSS (\u0026ordm;) were made in the lab. Twenty fruit samples were measured for length, width and thickness using a digital caliper with a precision of 0.01 mm. Additional morphological characteristics include Petiole length (mm), Petiole thickness (mm), Leaf area (cm2), and based on the leaf that was collected. The size, shape and volume of the fruit were described by the main morphometric parameters calculated from these measurements. A simple indicator of total fruit size, the Fruit Size Index (FSI) is computed as the product of fruit width and length (FSI\u0026thinsp;=\u0026thinsp;Length \u0026times; Width). The Fruit Shape Index measures how elongated a fruit is by dividing its length by its width (Shape Index\u0026thinsp;=\u0026thinsp;Length / Width). A round shape is suggested by values near 1, whereas elongated or flattened forms are indicated by values much greater or less than 1.\u003c/p\u003e\n \u003cp\u003eThe fruit\u0026apos;s degree of symmetry is expressed by the Fruit Roundness or Elongation Ratio (Roundness\u0026thinsp;=\u0026thinsp;Smaller dimension / Larger dimension \u0026times; 100), where higher percentages indicate more spherical fruits. Fruit Volume (approximate), the following formula was used to estimate fruit volume, assuming an ellipsoidal geometry.\u003c/p\u003e\n \u003cp\u003e\u003cimg src=\"https://myfiles.space/user_files/69519_bce2c0439cd956a6/69519_custom_files/img1773091019.png\"\u003e\u003c/p\u003e\n \u003cp\u003eWhere L is fruit length and W is fruit width.\u003c/p\u003e\n \u003cp\u003eThe surface area (approximate) was estimated using the ellipsoidal formula developed by Knud Thomsen.\u003c/p\u003e\n \u003cp\u003e\u003cimg src=\"https://myfiles.space/user_files/69519_bce2c0439cd956a6/69519_custom_files/img1773091046.png\"\u003e\u003c/p\u003e\n \u003cp\u003eWhere a\u0026thinsp;=\u0026thinsp;L/2, b\u0026thinsp;=\u0026thinsp;c = W/2, and p\u0026thinsp;\u0026asymp;\u0026thinsp;1.6.\u003c/p\u003e\n \u003cp\u003eCanopy volume for each tree was estimated with the \u0026ldquo;contour method\u0026rdquo; (Wright \u003cem\u003eet al.\u003c/em\u003e, 2005), treating the canopy as a geometric solid whose base widths \u0026lsquo;a\u0026rsquo; (perpendicular to the planting row) and \u0026lsquo;b\u0026rsquo; (parallel to the row) were measured at the canopy base, and whose height \u0026lsquo;h\u0026rsquo; was taken from the lowest branch to the apex. The contour functions \u0026lsquo;m(x)\u0026rsquo; and \u0026lsquo;m(y)\u0026rsquo; adjust for departures from an ideal canopy shape. The final expression is (CV = [(1\u0026frasl;4) \u0026pi; a b h) / (m(x)\u0026thinsp;+\u0026thinsp;m(y)\u0026thinsp;+\u0026thinsp;1].\u003c/p\u003e\n\u003c/div\u003e\n\u003cdiv id=\"Sec4\"\u003e\n \u003ch2\u003eData Analysis\u003c/h2\u003e\n \u003cp\u003eAll morphological characteristics of 30 wild Ber (\u003cem\u003eZiziphus spp\u003c/em\u003e.) accessions have been analysed for multivariate relationships between accessions. Principal component analysis (PCA) was performed to identify the traits that contribute the most to inter-genotypic variation, and a biplot of the first two PCs was used to visualise genotypic dispersion and trait loading. Hierarchical clustering has been calculated using Ward\u0026apos;s minimum variance algorithm (Ward\u0026apos;s D\u0026sup2;) for Euclidean distances. Pearson\u0026apos;s correlation coefficients were used to quantify pairwise trait associations, and a heatmap was created to illustrate both positive and negative relationships that are pertinent to selection. Python (v3.11) was used for all analyses, with Matplotlib/Seaborn for visualisation, SciPy for hierarchical clustering, and scikit-learn for PCA and scaling.\u003c/p\u003e\n\u003c/div\u003e"},{"header":"Results and Discussion","content":"\u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eCollection and survey results\u003c/h2\u003e \u003cp\u003eIn February and March 2019, systematic field surveys were conducted in the Kutch district of Gujarat to collect 30 geo-referencedber (\u003cem\u003eZiziphus spp.)\u003c/em\u003e accessions from the talukas of Bhuj, Nakhatrana, Mandvi, and Abdasa. The distribution of accessions was as follows: Mandvi (n\u0026thinsp;=\u0026thinsp;4; 13.33%), Bhuj (n\u0026thinsp;=\u0026thinsp;17; 56.67%), Nakhatrana (n\u0026thinsp;=\u0026thinsp;8; 26.67%), and Abdasa (n\u0026thinsp;=\u0026thinsp;1; 3.33%), with multiple sampling sites covered within each taluka (Table\u0026nbsp;1). The surveyed area spanned approximately 170 km, with the centroid located at 23.3688\u0026deg; N, 69.5494\u0026deg; E, extending from 23.0347\u0026ndash;23.6881\u0026deg; N and 69.2051\u0026ndash;70.7156\u0026deg; E. Three accessions represented the geographical extremes: Br_Mand_7 (southernmost; 23.0347\u0026deg; N, 69.3000\u0026deg; E), Br_Nakh_29 (northernmost and westernmost; 23.6881\u0026deg; N, 69.2051\u0026deg; E), and Br_Mand_8 (easternmost; 23.5629\u0026deg; N, 70.7156\u0026deg; E) (Fig.\u0026nbsp;1). Together, this coverage encompassed the major ber-growing tracts of central and western Kutch, ensuring adequate representation both within and across talukas for subsequent diversity analyses.Comparable botanical surveys in the Indian arid zone have similarly emphasized broad geographic coverage to capture genetic variability. For instance, (Kumar and Tripathi, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) documented 40 ber genotypes from eastern Uttar Pradesh (Dev et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) reported 27 genotypes of \u003cem\u003eGrewia tenax\u003c/em\u003e and 16 genotypes of \u003cem\u003eCordia gharaf\u003c/em\u003e from the Kachchh region of Gujarat; while (Venkatesan \u003cem\u003eet al\u003c/em\u003e., 2018) collected 40 \u003cem\u003eGrewia tenax\u003c/em\u003e genotypes from Barmer, Jaisalmer, Jodhpur, and Pali districts of western Rajasthan.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eDescriptive Statistics\u003c/h3\u003e\n\u003cp\u003eA wide range of phenotypic variation was observed in both fruit and plant architectural traits. Plant height varied from 144 to 455 cm, with an average of 255.25\u0026thinsp;\u0026plusmn;\u0026thinsp;77.49 cm (CV\u0026thinsp;=\u0026thinsp;30.36%). The tallest plant was Ber_Bhuj_30, while the shortest was Ber_Bhuj_1. Similar variability in plant height has been reported in other germplasm surveys; for example, Kumar and Tripathi (\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) documented heights ranging from 3.22 to 5.50 m across 40 genotypes, while Kumar et al., (\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) reported heights between 1.6 and 4.4 m. Significant differences were also recorded in plant spread. The north south spread ranged from 132 to 525 cm (CV\u0026thinsp;=\u0026thinsp;32.13%), whereas the east\u0026ndash;west spread varied between 147 and 540 cm (CV\u0026thinsp;=\u0026thinsp;36.66%), with Ber_Abda_20 showing the widest east\u0026ndash;west spread. Canopy volume exhibited the greatest variability, ranging from 0.58 to 8.34 m\u0026sup3; (CV\u0026thinsp;=\u0026thinsp;66.24%). The smallest canopy volume was observed in Ber_Bhuj_1 and the largest in Ber_Bhuj_30. Such wide variation in canopy traits has also been highlighted in other ber germplasm evaluations Kumar and Tripathi, (\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBranching habits showed substantial variation, with the number of primary branches per plant ranging from 2 to 20 (CV\u0026thinsp;=\u0026thinsp;62.19%). Leaf area also varied widely, from 1.38 to 9.76 cm\u0026sup2;. Leaf size is a key parameter influencing photosynthesis, as well as food, fodder, and biomass production. Comparable variation has been reported in \u003cem\u003eGrewia tenax\u003c/em\u003e germplasm from Kutch, Gujarat, where leaf size ranged from 3.1 to 8.3 cm\u0026sup2; (mean 4.8 cm\u0026sup2;; Dev et al., (\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Similar variability in leaf shape (cordate, oval, obovate and elliptic) and other morphological traits has also been documented in ber genotypes from eastern Uttar Pradesh (Kumar and Tripathi, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Such phenotypic diversity in morpho-physico-chemical traits has likewise been observed in other crops, as reported by \u0026Ouml;zrenk et al., (\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), Karadeniz et al., (\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2013\u003c/span\u003e); Dev et al., (\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). This diversity is likely influenced by multiple factors, including geographical distribution, origin, genotype, climatic conditions, and their complex interactions.\u003c/p\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eFruit Traits\u003c/h2\u003e \u003cp\u003eFor the selection of superior genotypes, fruit characteristics particularly fruit weight are among the most important target traits in ber improvement programs. In the present study, fruit size attributes varied from small to large types. The mean fruit length was 10.67 mm (range: 8.29\u0026ndash;13.05 mm; CV\u0026thinsp;=\u0026thinsp;10.14%) and the mean width was 11.27 mm (range: 8.77\u0026ndash;14.73 mm; CV\u0026thinsp;=\u0026thinsp;10.69%). The composite fruit size (length \u0026times; width) was smallest in Ber_Bhuj_2 and largest in Ber_Abda_20 (Table\u0026nbsp;2). Similar variability in fruit weight, 100‑seed weight, fruit length, and width has been documented in ber and other horticultural crops. For instance, such variation has been reported in ber (Kumar et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Sharif et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Meena et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), Grewiatenax (Dev et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), apple (Miller et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Kaya et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), fig (Darjazi, 2011), apricot (Kumar, 2015), and Cordia gharaf (Dev et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). In the present collection, fruit roundness, surface area, and fruit shape index were the most consistent attributes across genotypes.\u003c/p\u003e \u003cp\u003eFruit test weight (308\u0026ndash;1496 g), five‑fruit weight (1.54\u0026ndash;7.48 g), and fruit volume (353.20\u0026ndash;1332.10 mm\u0026sup3;) showed comparable variability (CV\u0026thinsp;\u0026asymp;\u0026thinsp;29\u0026ndash;32%), with the highest values consistently observed in Ber_Abda_20. Similar significant differences in fruit size and weight have also been reported in indigenous jujube cultivars of Pakistan, where the cultivar Foladi exhibited comparatively high fruit weight and width (Anjum et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Previous diversity studies have emphasized that variation in morphological traits provides a reliable basis for germplasm selection and improvement (Meghwal et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Kumar et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Dev et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Total soluble solids (TSS) displayed high variability (CV\u0026thinsp;=\u0026thinsp;37.99%), ranging from 3.40 to 17.50 \u0026deg;Brix, with Ber_Bhuj_21 recording the highest value. Similar variation (5.9\u0026ndash;13.2 \u0026deg;Brix) was also noted among wild ber genotypes in Pakistan (Sharif et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Among Indian cultivars, TSS values ranged more widely (12.5\u0026ndash;24.2 \u0026deg;Brix), which may be attributed to genetic differences, cultural practices, and stage of ripening Anjum et al., (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Consistent with this trend, cultivars such as Khobani (14.92 \u0026deg;Brix) and Dilbahar (11.33 \u0026deg;Brix) showed contrasting TSS levels, Anjum et al., (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e); Kumari et al. (\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) further reported that the cultivar Sanaur‑4 recorded high TSS (15.76 \u0026deg;Brix) and a superior TSS/acid ratio (40.38), along with distinct sugar profiles and vitamin C content across genotypes. In the current study, canopy volume, branching pattern, and stem girth were among the most variable traits, whereas fruit roundness and shape remained relatively stable across accessions. Based on a combination of vigour, fruit size, and quality attributes, the accessions Ber_Abda_20, Ber_Bhuj_21, Ber_Bhuj_29, and Ber_Bhuj_30 were identified as the most promising candidates for further evaluation and potential use in crop improvement programs.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003ePrincipal Component Analysis\u003c/h3\u003e\n\u003cp\u003ePrincipal component analysis (PCA) reduced the original 17 traits to four principal components (PCs) with eigenvalues greater than 1, together explaining 85.55% of the total variation. PC1 (eigenvalue\u0026thinsp;=\u0026thinsp;8.817; 50.14% variance) showed strong positive loadings for fruit width, fruit length, five‑fruit weight, size index, fruit volume, fruit surface area, test weight, and total soluble solids (TSS). This axis represented accessions combining large fruit size with improved quality traits, such as Ber_Abda_20 and Ber_Bhuj_30. PC2 (eigenvalue\u0026thinsp;=\u0026thinsp;3.605; 20.50% variance) contrasted vegetative vigour against fruit traits (Fig.\u0026nbsp;2). High positive loadings were recorded for plant height, canopy spread, stem girth, canopy volume, and branch number, whereas fruit length, width, size index, shape index, and roundness loaded negatively. This axis separated tall, vigorous types (e.g., Ber_Bhuj_30, Ber_Bhuj_21) from compact, fruit-dominant accessions. PC3 (eigenvalue\u0026thinsp;=\u0026thinsp;1.515; 8.62% variance) was associated primarily with fruit shape traits (shape index and roundness), while PC4 (eigenvalue\u0026thinsp;=\u0026thinsp;1.107; 6.29%) highlighted leaf area and stem girth (Fig.\u0026nbsp;3). Similar PCA findings in ber have shown that the first two PCs often capture a major share of variation, typically separating yield‑related traits from fruit quality attributes Anjum et al., (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e); Kumar et al., (\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) reported that the first two PCs explained 62.97% of total variation with one group characterized by superior yield traits and the other by fruit quality. Comparable patterns have also been described in rose germplasm, where fruit diameter, fruit weight, pulp weight, and seed number explained substantial proportions of the total variation (Verma, 2015).\u003c/p\u003e\n\u003ch3\u003eHierarchical Cluster Analysis\u003c/h3\u003e\n\u003cp\u003eUsing a Euclidean distance dendrogram, four distinct clusters were identified. Cluster 1 consisted of high-vigor, large-fruited types (Ber_Bhuj_21, Ber_Nakh_22, Ber_Bhuj_24, Ber_Bhuj_29, and Ber_Bhuj_30) characterized by tall plants, broad canopies, and large fruits. Cluster 2 included a single accession (Ber_Abda_20) that exhibited moderate vigor combined with extremely large fruits and high total soluble solids (TSS). Cluster 3 comprised eight compact accessions with low vigor and small, round fruits (e.g., Ber_Bhuj_1, Ber_Bhuj_2). Cluster 4 contained 16 accessions with intermediate vigour and fruit size (Fig.\u0026nbsp;4). Similar grouping patterns based on pomological traits have been reported in cluster analyses of indigenous Indian jujube Anjum et al., (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), Wild Ber Sharif et al., (\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), domesticated and wild Ber Sharif et al., (\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), and Wild Indian Ber Sareen \u003cem\u003eet al.\u003c/em\u003e, (2023).\u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eCorrelation Analysis\u003c/h2\u003e \u003cp\u003eThe correlation analysis revealed that plant height was strongly correlated with canopy volume (r\u0026thinsp;\u0026asymp;\u0026thinsp;0.91), east\u0026ndash;west spread (r\u0026thinsp;\u0026asymp;\u0026thinsp;0.66), and north\u0026ndash;south spread (r\u0026thinsp;\u0026asymp;\u0026thinsp;0.73). Fruit weight (measured from 5 fruits) showed positive correlations with fruit width (r\u0026thinsp;\u0026asymp;\u0026thinsp;0.47), size index (r\u0026thinsp;\u0026asymp;\u0026thinsp;0.58), volume (r\u0026thinsp;\u0026asymp;\u0026thinsp;0.61), surface area (r\u0026thinsp;\u0026asymp;\u0026thinsp;0.61), and test weight (r\u0026thinsp;\u0026asymp;\u0026thinsp;0.82). The shape index (length/width) was negatively correlated with width (r\u0026thinsp;\u0026asymp;\u0026thinsp;\u0026minus;\u0026thinsp;0.40), size index (r\u0026thinsp;\u0026asymp;\u0026thinsp;\u0026minus;\u0026thinsp;0.46), and surface area (r\u0026thinsp;\u0026asymp;\u0026thinsp;\u0026minus;\u0026thinsp;0.40). Total soluble solids (TSS) demonstrated moderate positive correlations with plant height, canopy spreads, and fruit dimensions (Fig.\u0026nbsp;5). Similar correlations have been reported in wild jujube populations, where leaf length and width showed a strong positive correlation (r\u0026thinsp;=\u0026thinsp;0.897), and fruit weight was negatively correlated with stone width (r\u0026thinsp;=\u0026thinsp;\u0026minus;\u0026thinsp;0.409) Sharif et al., (\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), supporting the present findings. Significant positive correlations were also observed for leaf dimensions, fruit weight, and fruit size (Azam-Ali et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2001\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e"},{"header":"Conclusion","content":"\u003cp\u003eThe study surveyed and collected 30 geo-referenced ber accessions from the Kutch district, revealing significant phenotypic and genetic diversity in plant and fruit traits. Multivariate analyses identified four distinct clusters, highlighting promising accessions like Ber_Abda_20 and Ber_Bhuj_30 for breeding. Strong correlations between vegetative growth and fruit quality traits were observed, providing valuable insight for future ber improvement and conservation in arid regions of Gujarat.\u003c/p\u003e"},{"header":"Declarations","content":" \u003cp\u003e \u003cb\u003eFunding Declaration\u003c/b\u003e \u003c/p\u003e \u003cp\u003eThis research did not receive any specific grant from funding agencies in the public, commercial, or not-for-profit sectors. All costs associated with the research and manuscript preparation were covered by the authors' institution.\u003c/p\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003ePermissions to collect the plants/plant parts\u003c/h2\u003e \u003cp\u003eThe plant materials used in this study were collected from non-protected areas and involved non-endangered, commonly cultivated species. The species investigated is a commonly cultivated crop, and no specific permits or licenses were required for collection.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eData Availability Statement\u003c/h2\u003e \u003cp\u003eThe datasets generated during and/or analysed during the current study are available from the corresponding author on reasonable request.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eEthics and Consent to Participate\u003c/h2\u003e \u003cp\u003eThe plant material used in this study was obtained from cultivated sources. All procedures involving plant collection complied with relevant local and national guidelines and regulations. Samples were collected from non-protected areas, and all relevant institutional and regulatory requirements were followed.\u003c/p\u003e \u003c/div\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eR.D. write the manuscriptT.S., C.K., A.T., D.D. collection and analyses of Data, All authors reviewed the manuscript.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAnjum MA, Rauf A, Bashir MA, Ahmad. The evaluation of biodiversity in some indigenous Indian jujube (\u003cem\u003eZiziphus mauritiana\u003c/em\u003e) germplasm through physio-chemical analysis. 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J Innovative Sci. 2022;8(1):97\u0026ndash;112. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.17582/journal.jis/8.1.97.112\u003c/span\u003e\u003cspan address=\"10.17582/journal.jis/8.1.97.112\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSharif N, Jaskani MJ, Naqvi SA, Awan FS. Exploitation of diversity in domesticated and wild ber (\u003cem\u003eZiziphus mauritiana\u003c/em\u003e Lam.) germplasm for conservation and breeding in Pakistan. Sci Hort. 2019;249:228\u0026ndash;39.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSrivastava AK, Singh SP, Singh HK. Seed variability in ber (\u003cem\u003eZiziphus\u003c/em\u003e species) collected from different localities of India. Indian J Plant Genetic Resour. 2001;14(2):176\u0026ndash;80.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eVenkatesan K, Patidar A, Singh M, Kumar M, Kumawat RN, Dev R, Uchoi J, Kumar A, Mertia DS, Singh JP. Distribution, associated vegetation, conservation and utilization of \u003cem\u003eGrewia tenax\u003c/em\u003e: an important underutilized shrub species of the Thar Desert of India. Plant Genetic Resour. 2019;17(1):73\u0026ndash;80.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eVerma MK, Lal S, Ahmed N, Kumar D, Singh DB, Sagoo PA. Genetic diversity among native wild hip rose (\u003cem\u003eRosa canina\u003c/em\u003e L.) genotypes collected from Kashmir valley. Indian J Hortic. 2015;72(2):250\u0026ndash;6.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWright H, Nichols D, Embree C. Evaluating the accountability of trunk size and canopy volume models for determining apple tree production potential across diverse management regimes. Acta Hort. 2006;707:237\u0026ndash;43. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://dx.doi.org/10.17660/ActaHortic.2006.707.30\u003c/span\u003e\u003cspan address=\"10.17660/ActaHortic.2006.707.30\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e.\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003eTables are available in the Supplementary Files section.\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"discover-plants","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"","sideBox":"Learn more about [Discover Plants](https://link.springer.com/journal/44372)","snPcode":"44372","submissionUrl":"https://submission.springernature.com/new-submission/44372/3","title":"Discover Plants","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Discover Series","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"Ziziphus mauritiana, germplasm diversity, Kutch district, morphological traits, principal component analysis, cluster analysis, fruit quality.","lastPublishedDoi":"10.21203/rs.3.rs-8618673/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8618673/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThis study assessed morphological diversity in ber (\u003cem\u003eZiziphus spp.\u003c/em\u003e) germplasm collected from Gujarat\u0026rsquo;s Kutch district. Thirty accessions from Bhuj, Nakhatrana, Mandvi, and Abdasa talukas were sampled across ~\u0026thinsp;170 km, capturing the central and western Kutch ber-growingarea. Seventeen morphological and quality traits including plant height, canopy spread, branch number, leaf area, fruit dimensions, weight and total soluble solids (TSS) were measured. Descriptive statistics showed broad phenotypic dispersion: plant height ranged from 144 to 455 cm; canopy volume varied from 0.58\u0026ndash;8.34 m\u0026sup3;; and leaf area from 1.38\u0026ndash;9.76 cm\u0026sup2;, TSS ranged from 3.40\u0026ndash;17.50 \u0026deg;Brix. Fruit length, width, and 5-fruit weight displayed moderate variability. Principal component analysis reduced the 17 traits to four components, explaining 85.55% of total variation. PC1 (50.14%) was associated with fruit dimensions and TSS, identifying high-yield, high-quality accessions (e.g., Ber_Abda_20, Ber_Bhuj_30); PC2 (20.50%) contrasted vegetative vigour with fruit size. Hierarchical clustering separated accessions into four clusters: (1) tall, large-fruited types; (2) a unique accession combining moderate vigour and very large, sweet fruits; (3) compact, small-fruited types; and (4) intermediate vigour and fruit size. Correlation analysis revealed strong positive associations between plant height and canopy volume (r\u0026thinsp;\u0026asymp;\u0026thinsp;0.91) and between fruit weight and size index (r\u0026thinsp;\u0026asymp;\u0026thinsp;0.58), while TSS correlated moderately with plant size and fruit dimensions. Overall, the Kutch ber germplasm displayed considerable morphological and biochemical diversity. The promising accessions Ber_Abda_20, Ber_Bhuj_21, Ber_Bhuj_29 and Ber_Bhuj_30 offer potential for cultivar development and highlight the value of conserving Kutch\u0026rsquo;s under-studied genetic resources.\u003c/p\u003e","manuscriptTitle":"Accessing agro-morphological variation in Ber (Ziziphus spp.) Germplasm from Kutch, Gujarat, Using Multivariate Techniques","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-03-09 21:20:25","doi":"10.21203/rs.3.rs-8618673/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"editorInvitedReview","content":"","date":"2026-03-23T03:38:48+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-03-22T03:40:02+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-03-16T14:49:47+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-03-12T05:39:19+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"122378853213511615935197427325299282102","date":"2026-03-11T04:46:03+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"48939410371656804469312303154434805685","date":"2026-03-11T02:53:22+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-03-10T11:12:16+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"250877004518688834362861407744357397563","date":"2026-03-09T11:19:42+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"171080818870454533018135647910003500421","date":"2026-03-09T11:05:01+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"267417519162752212272760545630630970829","date":"2026-03-09T10:52:53+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"75972574441087551606918582420544464780","date":"2026-03-06T10:59:36+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"61791806561238204128371750290013198791","date":"2026-03-05T00:16:40+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-03-04T10:33:48+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"","date":"2026-02-13T10:32:22+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-02-11T06:23:01+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2026-02-06T11:05:18+00:00","index":"","fulltext":""},{"type":"submitted","content":"Discover Plants","date":"2026-02-06T10:13:15+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"discover-plants","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"","sideBox":"Learn more about [Discover Plants](https://link.springer.com/journal/44372)","snPcode":"44372","submissionUrl":"https://submission.springernature.com/new-submission/44372/3","title":"Discover Plants","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Discover Series","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"7906f7ef-f385-4783-9704-e3dc73bb27f2","owner":[],"postedDate":"March 9th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-03-09T21:20:25+00:00","versionOfRecord":[],"versionCreatedAt":"2026-03-09 21:20:25","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8618673","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8618673","identity":"rs-8618673","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}