Trait diversity in wild and parkland baobab (Adansonia digitata L.) populations in Malawi

Trait diversity in wild and parkland baobab (Adansonia digitata L.) populations in Malawi | 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 Trait diversity in wild and parkland baobab (Adansonia digitata L.) populations in Malawi Herbert Jenya, Owen Kachala This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9317216/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 11 You are reading this latest preprint version Abstract Baobab ( Adansonia digitata L.) is a multipurpose fruit tree with high nutritional and economic importance across sub-Saharan Africa. We characterised fruit, seed, and tree trait diversity in five wild and parkland baobab populations in Malawi to generate baseline information for conservation and germplasm selection. A total of 135 trees were sampled. Thirteen fruit shapes were identified, with ellipsoid the most common across all populations. Eleven quantitative fruit and seed traits differed significantly among the 13 fruit shapes (p ≤ 0.001). High spheroid fruits consistently showed superior productivity, with the greatest fruit weight (235 ± 17 g), fruit width (9.6 ± 0.3 cm), pulp weight (45.3 ± 3.9 g), total seed weight (98.1 ± 8.1 g), and number of seeds per fruit (208 ± 16). Oblong fruits ranked second for most weight-related traits. PCA identified two principal axes explaining 66.4% of the total variance. PC1 (42.0%) represented a fruit productivity gradient associated with fruit weight, seed weight, and seed number, whereas PC2 (24.4%) captured independent variation in seed size traits, including length, width, and thickness. Diameter at breast height (DBH) also varied significantly among populations (F = 12.25, p < 0.001), with Karonga supporting the largest trees and Neno the smallest. Overall, the substantial trait diversity documented, particularly in Karonga and Salima, provides a strong foundation for conservation, domestication, and selection of superior baobab germplasm in Malawi. Conservation Domestication and management Farmland Morphology Selection Superior Germplasm Ideotypes Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 1.0 INTRODUCTION Adansonia digitata L. belongs to the subfamily Bombacoideae within the family Malvaceae. The genus Adansonia comprises eight species, of which A. digitata is the widespread African species (Wickens & Lowe, 2008 ). Commonly known as baobab, monkey bread tree, or bottle tree, it is one of Africa’s most iconic long-lived tree species and is valued for its ecological, cultural, and economic importance (Kehlenbeck et al. 2015 ; Sidibé and Williams 2002 ; Wickens and Lowe 2008 ). The tree is characterized by massive trunks, broad spreading branches, and a large rounded canopy (Sidibé and Williams 2002 ). Some individuals are exceptionally old, with ages extending well beyond a millennium (Patrut et al. 2018 ). Over the past two decades, increasing research attention has positioned baobab as an important orphan tree species for food and nutrition security, livelihood support, and agroforestry development (Assogbadjo et al. 2021 ). In Malawi, baobab occurs mainly in wild stands in savannah landscapes and on farmland. Baobab remains important in local diets and as a source of income for rural communities (Gebauer and Luedeling 2013 ; Kamatou et al. 2011 ). Numerous products are made locally, including baobab pulp, sweets, ice lollies, juice, jam, coffee substitutes, soap, and lip balm (Munthali 2012 ). Additionally, products in wider markets include beverages, cereals, cereal bars, probiotic yogurts, and cosmetic products derived from baobab seed oil (Gruenwald 2009 ; Gruenwald and Galizia 2005 ; Kamatou et al. 2011 ). Different plant parts, including the roots, bark, leaves, and seeds, are also used in traditional medicine to treat various ailments (Buchmann et al. 2010 ; Wickens and Lowe 2008 ). Recent studies further show that baobab fruit pulp is rich in key nutrients and that its composition varies geographically, reinforcing the species’ growing importance in food and nutrition research as well as the need to identify superior germplasm (Assogbadjo et al. 2021 ; Stadlmayr et al. 2020 ). The commercialization of baobab products continues to contribute to rural livelihoods in areas where the species is abundant. Recent studies show that its value now extends beyond household use to increasingly organized local and regional value chains. In Malawi, the baobab sector has in recent years shifted from mainly subsistence use toward wider business opportunity, although this transition remains constrained by weak policy support, rising quality requirements, and uneven value chain coordination (Meinhold et al. 2022). Recent evidence from Tanzania likewise shows that baobab fruit value chains generate economic value through margins, wages, and taxes, but are limited by poor storage, low prices, lack of capital, poor processing facilities, seasonal fruit supply, and limited access to market information (Ngole et al. 2024 ). Studies from Malawi further show that postharvest losses reduce the value ultimately captured by collectors, traders, and processors, highlighting inefficiencies in the chain (Cossam et al. 2023 ). These newer findings complement earlier evidence that baobab trade contributes meaningfully to household livelihoods in southern Africa (Luckert et al. 2014 ). This implies that the long-term economic potential of baobab will depend on stronger value chain organization, reduced losses, better processing and storage, and more deliberate domestication and management of the resource base. Conservation concerns have become increasingly important in recent years. Emerging evidence suggests that baobab populations in several parts of Africa are under pressure from land-use change, agricultural expansion, climate variability, and weak natural regeneration (Abere et al. 2023 ; Assogba et al. 2022 ; Venter and Witkowski 2010 ). Studies on habitat suitability under present and future climates indicate that persistence of the species will depend not only on protecting existing stands, but also on supporting regeneration, restoration, and cultivation within farming systems (Assogba et al. 2022 ). In addition, recent syntheses emphasize that, despite growing interest in baobab, the species remains underexploited relative to its potential and still faces important biological and management constraints that could limit its contribution to food systems and rural livelihoods if conservation and domestication efforts are not strengthened (Assogbadjo et al. 2021 ). Knowledge of trait variation in baobab is crucial for selecting plant material with desirable characteristics for domestication, conservation, and improvement (Santos et al., 2012 ). Previous studies have documented phenotypic variation in baobab fruits, leaves, seeds, and tree traits in Malawi and elsewhere in Africa (Assogbadjo et al. 2005 , 2006 , 2011 ; Cuni Sanchez et al. 2010 , 2011 ; De Smedt et al. 2011 ; Munthali et al. 2012 ; Omondi et al. 2019 ). More recent work also shows substantial genetic diversity and population structure in A. digitata , strengthening the case for conserving diverse germplasm while identifying superior material for domestication (Chládová et al. 2019 ). Likewise, evidence of early genetic variation in survival and growth traits indicates that baobab improvement and breeding are feasible and can support future domestication efforts (Kalinganire et al. 2023 ). Fruit shape may be an especially important trait because it may influence consumer preference, marketability, productivity, and the development of ideotypes for domestication (Rashidi and Seyfi 2007; Vihotogbé et al. 2013). Multivariate approaches such as principal component analysis and cluster analysis have proven useful in identifying independent axes of phenotypic variation and grouping baobab populations for domestication prioritisation (Assogbadjo et al. 2006 ; De Smedt et al. 2011 ). Tree morphological characteristics, including diameter at breast height (DBH), also vary among populations and may reflect differences in population age structure, management history, and land-use context (Assogbadjo et al. 2006 ; Venter and Witkowski 2010 ). However, important gaps remain for Malawi. Although previous studies have examined fruit and seed traits in selected wild populations (Munthali et al. 2012 ), information is still limited on the diversity of fruit shapes across populations, the relationship between fruit shape and quantitative fruit and seed traits, and the extent to which fruit productivity and seed morphology represent independent axes of variation. An integrated characterisation that combines fruit shape diversity indices, population-level trait variation, multivariate structure, and tree morphological attributes is therefore needed to support conservation, domestication, and selection of superior germplasm in Malawian baobab populations. This study aimed to characterise the trait diversity of baobab across five wild and parkland populations in Malawi to inform conservation, management, improvement, and domestication efforts. Specifically, it addressed the following questions: (1) What fruit shapes occur, and do they differ consistently between individual trees and among populations? (2) Do quantitative fruit traits vary among fruit shapes? (3) Do quantitative and qualitative seed traits vary among fruit shapes? (4) How does fruit shape diversity, quantified using the Shannon–Wiener diversity index, compare across the five populations? (5) Do fruit and seed traits differ significantly among populations? (6) What is the multivariate structure of fruit and seed phenotypic variation, and are fruit productivity and seed morphology independent axes of variation? and (7) How do tree morphological characteristics and diameter at breast height vary among populations? We hypothesize that baobab populations in Malawi exhibit significant phenotypic variation in fruit shape, fruit traits, seed traits, and tree morphology. Specifically, fruit-shape categories are expected to differ in associated quantitative fruit and seed characteristics, populations are expected to differ in fruit-shape diversity and overall trait composition, and multivariate analysis is expected to reveal distinct dimensions of variation separating fruit productivity traits from seed morphology traits. 2.0 MATERIALS AND METHODS 2.1 Study site Fruit samples were collected from five baobab populations in Chikwawa, Karonga, Mangochi, Neno, and Salima (Table 1 ). Study populations were selected to represent the major agro-ecological settings across Malawi in which the species occurs, based on national agro-ecological classifications reported by Government of Malawi (Government of the Republic of Malawi 2005 ). The sites span considerable variation in altitude, mean annual rainfall (MAR), temperature, and soil type (Table 1 ). Chikwawa, situated in the Lower Shire Valley, is characterised by low elevation (< 200 m), MAR of 710–850 mm, mean annual temperatures exceeding 25°C, and predominantly vertisol soils. Mangochi and Salima, both located in the lakeshore plains, receive similar rainfall (710–850 mm) with alluvial calcimorphic soils overlying vertisols and mean annual temperatures of 20–25°C. Neno, in the southern highlands, receives 1,200–1,600 mm of rainfall annually with sandy ferralitic soils and cooler temperatures of 19–21°C. Karonga, in the northern lakeshore zone, is the wettest site (MAR > 1,600 mm) with predominantly ferrosols and regosols and mean annual temperatures of 23–25°C (Table 1 ). Further, populations were selected based on abundance of trees and prevailing trade in baobab products. Table 1 Location, physical description of baobab populations and number of trees sampled in Malawi Population Coordinates Agro-ecological zone Altitude Mean annual rainfall (mm) Mean annual temperature Soil type Chikwawa (21**) 16°02’S, 34°50’E Shire Valley 25 Vertisols Mangochi (30**) 14°25’S, 35°16’E Lakeshore plains 200–1200 710–850 20–25 Alluvial calcimorphic soils above the vertisols Salima (30**) 13°47’S, 34°28’E Lakeshore plains 200–1200 710–850 20–25 Alluvial calcimorphic soils above the vertisols Neno (30**) 15°25’S, 34°40’E Southern Highlands 900–1500 1200–1600 19–21 Sandy ferralitic Karonga (24**) 09°56ˈS 33 ° 56ˈE Northern Lakeshore 475–1000 > 1600 23–25 Ferrosols dominant; regosols ** number of trees sampled 2.2 Data collection Fruits were collected at the peak of fruit maturity between March and May of 2016. Dry fruits (10 per tree) were randomly picked from trees. A total of 135 trees were characterized from the five populations (Table 1 ). Fruit shape description was done according to Gurashi and Kordofani ( 2014 ) and Kehlenbeck et al. ( 2015 ). From each fruit shape, subsamples of five fruits were randomly selected for the assessment of fruit, seed, and pulp weight. Fruits were weighed to the nearest 1 g with an electronic scale (TREE LW Measurements HRB 10001-High Resolution Balance 10,000 g ×0.1 g). Fruit shells were cracked manually and the fruit content (pulp + seeds) were removed and weighed moist. Pulp was separated from seeds by washing with water, and seeds were then air dried and weighed again. Pulp content was calculated by subtracting the moist seed weight from the fruit content (pulp + seeds). Seeds were counted to determine the seed number per fruit. Single seed weight was calculated by dividing the seed weight by the number of seeds according to Cuni Sanchez et al., ( 2011 ). A subsample of ten seeds per fruit was randomly selected for the measurement of seed length, seed width, and seed thickness using a micro-calliper to the nearest mm. Seed shape and colour were visually assessed in the laboratory using the guide proposed by Kehlenbeck et al. ( 2015 ). Tree morphological traits were recorded for each of the 135 sampled trees at the time of fruit collection. Diameter at breast height (DBH) was measured at 1.3 m above ground level using a diameter tape. Crown shape, trunk shape, bark colour, and bark texture were assessed visually for each tree following the descriptors proposed by Kehlenbeck et al. ( 2015 ). Crown shape was classified as roundish, ellipsoid, or semi-circular. Trunk shape was categorised as cylindrical, bell-shaped, concave, or cone-shaped. Bark colour was recorded as grey or reddish-brown, and bark texture as smooth or rough. The branch growth habit of each tree was recorded as erect, spreading, or drooping. The land use context of each tree was also noted, classifying trees as occurring in parkland, fallow land, forest/woodland, backyard garden, or shrubland. 2.3 Statistical analysis Fruit shape occurrence was analysed using descriptive statistics. Quantitative fruit and seed traits were compared among fruit shapes and among populations (districts) using one-way ANOVA with Sidak pairwise comparisons and compact letter displays were generated using the agricolae package (de Mendiburu 2023 ) in R (R Core Team 2024). Principal Component Analysis (PCA) was performed on standardised quantitative traits to examine the multivariate structure of variation among populations, using the FactoMineR package (Lê et al. 2008 ). Fruit shape diversity within each population was quantified using the Shannon-Wiener diversity index (H' = −Σ p i ln p i ), where p i is the relative frequency of each fruit shape, and Pielou's evenness (J' = H'/ln S, S = number of shapes), computed using the vegan package (Oksanen et al. 2024). Tree morphological traits were summarised using frequency distributions by district. Diameter at breast height (DBH) was compared among districts using one-way ANOVA with Sidak pairwise comparisons. All statistical analyses were conducted in R version 4.5.2 (R Core Team 2024). 3.0 RESULTS 3.1 Fruit Shape Diversity among Populations The Shannon–Wiener diversity index was highest in Chikwawa and Neno, which supported the highest fruit shape richness and diversity (H' = 2.30, S = 10 shapes each), followed by Karonga (H' = 2.08, S = 8). Mangochi and Salima had the fewest shapes (H' = 1.95, S = 7). Pielou's evenness was J' = 1.00 in all populations, a reflection of the balanced sampling design rather than a biological pattern (Table 2 ). Table 2 Fruit shape diversity indices for five baobab populations in Malawi. Population Shannon (H') No. of shapes (S) Evenness (J') Chikwawa 2.30 10 1 Karonga 2.08 8 1 Mangochi 1.95 7 1 Neno 2.30 10 1 Salima 1.95 7 1 Note: Evenness (J' = 1) reflects balanced sampling design This study identified thirteen distinct fruit shapes across the sampled baobab populations, namely ellipsoid, oblong, ovate, fusiform, high spheroid, globose, ellipsoid pointed, oblong compressed, oblong pointed, rhomboid, spheroid emarginate, and clavate (Table 3 ; Fig. 1 ). Ellipsoid, oblong, and ovate were common to all five populations, while spheroid emarginate was unique to Mangochi (Table 2 ). Table 3 Fruit shape occurrence (%) in five baobab populations in Malawi Fruit shape Chikwawa (A) Karonga (L) Mangochi (Ba) Neno (J) Salima (Ba) South North South South Central Clavate 10 0 0 6 0 Ellipsoid 25 16 38 30 43 Ellipsoid pointed —0 3 7 3 3 Fusiform 15 —0 10 13 —0 High spheroid — 16 —0 6 10 Oblong 15 12 21 6 24 Oblong compressed —0 —0 —0 6 7 Oblong pointed 5 —0 —0 6 —0 Obovate 10 10 14 3 —0 Ovate 10 19 7 24 10 Rhomboid 5 12 —0 —0 —0 Spheroid 5 12 —0 —0 3 Spheroid emarginate —0 —0 3 —0 —0 3.2 Variation of Fruit and Seed Traits among Fruit Shapes Eleven fruit and seed traits measured differed significantly ( P ≤ 0.001) among the 13 fruit shapes (Table 4 ). High spheroid fruits were consistently the largest by weight-related measures heaviest (235 ± 17 g), widest (9.6 ± 0.3 cm), highest pulp content (45.3 ± 3.9 g), greatest seed mass (98.1 ± 8.1 g), and most seeds per fruit (208 ± 16) differing significantly from fusiform and ovate fruits in most traits. Oblong fruits ranked second for most weight metrics. In contrast, fusiform (119 ± 17 g) and ovate (131 ± 15 g) fruits were the lightest, narrowest, and had the fewest seeds. Fruit length followed a different pattern: oblong pointed (19.0 ± 0.9 cm), ellipsoid pointed (18.2 ± 0.6 cm), and clavate (17.6 ± 0.9 cm) were the longest, while spheroid (10.5 ± 0.7 cm) and spheroid emarginate (10.4 ± 1.3 cm) were the shortest. At the individual seed level, clavate fruits consistently produced the largest seeds across all four dimensions heaviest single seeds (0.59 ± 0.03 g), longest (12.4 ± 0.2 mm), widest (9.6 ± 0.2 mm), and thickest (7.4 ± 0.1 mm) while ovate and spheroid fruits consistently had the smallest seeds. Seed dimensions showed notably less variation among shapes than fruit-level traits (CVs 6–22% vs. 19–52%). Full means and Sidak groupings for all eleven traits are presented in Table 4 . Table 4 Variation of fruit and seed traits (mean ± SE) among the 13 fruit shapes in five baobab populations in Malawi. Means in the same column sharing the same letter do not differ significantly (Sidak, p < 0.05). FW = fruit weight, FL = fruit length, FWd = fruit width, PW = pulp weight, % Pulp = % pulp to fruit weight, TSW = total seed weight, NS = number of seeds, SSW = single seed weight, SL = seed length, SWd = seed width, ST = seed thickness. Fruit shape Fruit traits Seed traits FW (g) FL (cm) FWd (cm) PW (g) % Pulp TSW (g) NS SSW (g) SL (mm) SWd (mm) ST (mm) Clavate 168.4 ± 20.9abc 17.6 ± 0.9d 6.7 ± 0.4ab 27.0 ± 4.8ab 15.52 ± 1.53a 59.3 ± 10.0abc 99 ± 19ab 0.59 ± 0.03b 12.4 ± 0.2d 9.6 ± 0.2c 7.4 ± 0.1b Ellipsoid 182.4 ± 13.2abc 16.7 ± 0.6cd 8.1 ± 0.2bc 33.0 ± 3.0ab 17.90 ± 0.97a 68.6 ± 6.3abc 142 ± 12abc 0.50 ± 0.02ab 11.8 ± 0.1abcd 8.8 ± 0.1ab 7.0 ± 0.1ab Ellipsoid pointed 187.4 ± 13.2abc 18.2 ± 0.6d 8.5 ± 0.2cd 32.0 ± 3.0ab 17.07 ± 0.97a 69.4 ± 6.3abc 129 ± 12ab 0.54 ± 0.02b 12.1 ± 0.1cd 9.0 ± 0.1bc 7.1 ± 0.1ab Fusiform 118.7 ± 17.1a 17.2 ± 0.7d 6.2 ± 0.3a 24.7 ± 3.9a 19.34 ± 1.25a 42.7 ± 8.1a 91 ± 16a 0.47 ± 0.03ab 11.7 ± 0.2abcd 8.8 ± 0.2abc 6.7 ± 0.1a High spheroid 235.2 ± 17.1c 13.5 ± 0.7ab 9.6 ± 0.3d 45.3 ± 3.9b 18.78 ± 1.25a 98.1 ± 8.1c 208 ± 16c 0.48 ± 0.03ab 11.4 ± 0.2abc 8.7 ± 0.2ab 6.9 ± 0.1ab Oblong 200.5 ± 13.2bc 17.7 ± 0.6d 7.9 ± 0.2bc 37.2 ± 3.0ab 18.36 ± 0.97a 83.4 ± 6.3bc 163 ± 12bc 0.52 ± 0.02ab 11.9 ± 0.1bcd 9.1 ± 0.1bc 7.0 ± 0.1ab Oblong compressed 162.5 ± 20.9abc 17.6 ± 0.9d 7.5 ± 0.4abc 24.1 ± 4.8ab 14.70 ± 1.53a 67.8 ± 10.0abc 135 ± 19abc 0.51 ± 0.03ab 11.5 ± 0.2abcd 9.3 ± 0.2bc 6.9 ± 0.1ab Oblong pointed 160.8 ± 20.9abc 19.0 ± 0.9d 6.9 ± 0.4ab 31.4 ± 4.8ab 19.39 ± 1.53a 58.8 ± 10.0abc 124 ± 19abc 0.48 ± 0.03ab 11.3 ± 0.2abcd 8.6 ± 0.2ab 6.7 ± 0.1a Obovate 157.3 ± 13.2ab 12.1 ± 0.6ab 8.7 ± 0.2cd 25.4 ± 3.0a 16.56 ± 0.97a 64.0 ± 6.3abc 121 ± 12ab 0.53 ± 0.02ab 11.7 ± 0.1abcd 8.8 ± 0.1ab 7.1 ± 0.1ab Ovate 131.4 ± 14.8a 13.8 ± 0.6bc 7.8 ± 0.3bc 24.8 ± 3.4a 18.10 ± 1.08a 46.8 ± 7.0a 107 ± 14ab 0.43 ± 0.02a 11.1 ± 0.2a 8.2 ± 0.1a 6.9 ± 0.1ab Rhomboid 187.9 ± 20.9abc 15.0 ± 0.9bcd 9.1 ± 0.4cd 29.0 ± 4.8ab 15.54 ± 1.53a 73.2 ± 10.0abc 134 ± 19abc 0.55 ± 0.03ab 12.4 ± 0.2cd 8.8 ± 0.2abc 7.2 ± 0.1ab Spheroid 135.1 ± 17.1ab 10.5 ± 0.7a 8.2 ± 0.3bcd 25.5 ± 3.9a 18.80 ± 1.25a 57.5 ± 8.1ab 124 ± 16ab 0.45 ± 0.03ab 11.1 ± 0.2ab 8.2 ± 0.2a 6.8 ± 0.1a Spheroid emarginate 156.8 ± 29.6abc 10.4 ± 1.3ab 9.5 ± 0.6cd 22.0 ± 6.7ab 14.07 ± 2.17a 71.8 ± 14.1abc 135 ± 27abc 0.58 ± 0.05ab 11.8 ± 0.3abcd 9.4 ± 0.3bc 7.0 ± 0.2ab G/mean 170.2 15.5 8.1 30.2 17.55 66.6 133 0.50 11.7 8.8 7.0 Four seed colours (Fig. 2 ) were found among the fruit shapes, including light brown (16% of all seeds), brown (46%), dark brown (35%), and greyish brown (3%). Three seed shapes were found (Fig. 3 ). The predominant shape was reniform (55%) followed by very reniform (44%). The least shape was oblong seed (1%). All the seeds had spots in varying density on their seed coats. Light brown seed (A), dark brown seed (C) and greyish brown seed (D) were light spotted while brown seed (C) were densely spotted. 3.5 Principal Component Analysis of Fruit and Seed Traits Principal Component Analysis of the 11 standardised quantitative traits yielded three principal components (PCs) with eigenvalues > 1, together explaining 77.3% of total variance. PC1 explained 42.0% of variance and was strongly loaded by overall fruit size traits: fruit weight (r = 0.967), seed weight (r = 0.941), pulp weight (r = 0.861), number of seeds (r = 0.825), and fruit width (r = 0.793), and thus represents a latent fruit productivity axis. PC2 explained 24.4% of variance and was dominated by individual seed dimension traits seed length (r = 0.802), seed thickness (r = 0.742), seed width (r = 0.739), and single seed weight (r = 0.724) representing a seed size axis. PC3 explained 10.8% of variance and was primarily associated with pulp percentage (r = 0.816) and fruit length (r = 0.380), representing a pulp quality axis. The biplot (Fig. 4 ) showed partial separation of populations along PC1, with Karonga and Salima positioned towards the high fruit productivity end, while Chikwawa clustered at the low end. Populations overlapped considerably along PC2, indicating broadly similar individual seed dimensions across populations. Neno and Chikwawa showed wider 90% confidence ellipses, indicating greater within-population variability in overall fruit traits. 3.6 Tree Morphological Characteristics and Diameter at Breast Height Bark colour was predominantly grey across all five districts (93.7% of trees overall), with reddish-brown bark recorded mainly in Karonga (26.7%). The most common tree shape was roundish (55.6% of all trees), followed by semi-circular (33.3%) and ellipsoid (11.1%). Mangochi was distinguished by a dominance of semi-circular canopy form (93.3%), while all sampled trees in Chikwawa were roundish (100%). Cone-shaped trunks were most prevalent across all districts (55.6%), followed by cylindrical (21.4%), concave (15.9%), and bell-shaped (7.1%). Spreading growth habit was most common overall (65.1%), though Chikwawa had a notably higher proportion of drooping trees (61.9%), and erect growth was most prevalent in Salima (26.7%) and Neno (23.3%). Trees were predominantly collected from fallow land, except in Chikwawa (95.2% on farmland) and Mangochi (50% in forest/woodland), suggesting differences in land-use and conservation context across populations (Fig. 8). DBH varied significantly among districts (F = 12.25, p < 0.001; Table 5 ). Karonga had the largest trees (mean 309.2 ± 16.6 cm), significantly larger than Neno (187.0 ± 7.9 cm) and Mangochi (227.0 ± 9.1 cm; both p < 0.001). Chikwawa (272.8 ± 10.3 cm) and Salima (231.2 ± 9.4 cm) were intermediate. The overall DBH range across all trees was 104–422 cm (Table 7). Table 5 Diameter at breast height (DBH) of baobab trees across five districts in Malawi. Different letters indicate significant differences (Tukey HSD, p < 0.05). District n (trees) Mean ± SE (cm) Range (cm) Tukey Chikwawa 21 273.1 ± 10.7 196.1–347 ab Karonga 15 309 ± 19.5 179.9–422 a Mangochi 30 226.7 ± 10.9 133.1–356.5 bc Neno 30 186.9 ± 9.5 120–294.4 c Salima 30 231.2 ± 12.9 104.1–374.6 b F = 12.25, p < 0.001. Different letters indicate significant differences (Tukey HSD, p < 0.05). 4.0 Discussion This study demonstrates substantial phenotypic variation in fruit and seed traits among wild and parkland baobab populations in Malawi, highlighting important opportunities for domestication, germplasm selection, and conservation. These differences are not merely descriptive; they point to functionally important diversity relevant to agroforestry systems where baobab contributes food, income, and resilience. This supports earlier work identifying Adansonia digitata as a nutritionally important but still under-domesticated tree species for which trait characterization remains a necessary step toward improvement and wider use (Kehlenbeck et al. 2015 ; Assogba et al. 2022 ; Kitony et al. 2024 ). The occurrence of 13 fruit-shape classes across five populations indicates considerable intraspecific diversity. This exceeds some earlier descriptions from southern Africa and aligns more closely with later studies elsewhere in Africa (Wickens 1982; Gebauer and Luedeling 2013 ; Gurashi and Kordofani, 2014 ). The fact that some fruit shapes observed in Malawi were not reported in Sudan further suggests that baobab morphological diversity remains incompletely documented across its range. This is important because it indicates that southern African populations should not be viewed as morphologically marginal, but rather as potentially valuable reservoirs of undercharacterized diversity. A key finding was that each sampled tree consistently produced a single fruit-shape type, with no within-tree polymorphism. Although the present study did not test heritability directly, this within-tree consistency suggests that fruit shape is a relatively stable phenotype and may be under substantial genetic or maternal developmental control. For agroforestry domestication, this is encouraging because it strengthens the rationale for selecting elite mother trees and propagating them vegetatively, consistent with earlier baobab domestication studies and successful grafting work in Malawi and Kenya (Tchoundjeu et al. 2006; Anjarwalla et al. 2017 ; Jenya et al. 2018 ). Recent genomic evidence showing substantial population structure and adaptive diversity in baobab further supports the view that at least part of the observed variation has a biological basis (Kitony et al. 2024 ). Fruit shape was also associated with clear differences in productivity. Variation in fruit weight, pulp weight, and total seed weight shows that fruit shape is not simply a descriptive character but one linked to traits of practical value. Similar patterns have been reported in Sudan and Mali (De Smedt et al. 2011 ; Gurashi and Kordofani, 2014 ). In this study, high spheroid and oblong fruits appear particularly promising where the objective is to maximize pulp and total seed output. However, the results also show that productivity is multidimensional. Clavate fruits produced the largest individual seeds, whereas high spheroid fruits tended to maximize total seed mass through greater seed number. This suggests that selection for pulp-oriented production and selection for seed-oriented uses may favor different phenotypes. That distinction matters in agroforestry settings, where baobab may be managed for multiple products including pulp, oil, propagation material, and nursery stock. A single ideotype is therefore unlikely to serve all production goals. The identification of four seed-coat colour classes further suggests that current baobab descriptor systems may not yet capture the full phenotypic range present in eastern and southern Africa (Kehlenbeck et al. 2015 ). This could have practical relevance because seed-coat colour in other species has been linked to biochemical composition and dormancy-related traits (Dixon and Sumner 2003 ; Ochuodho and Modi 2008 ). Although such relationships were not tested here, the finding points to useful follow-up work on germination and nutritional quality. By contrast, seed shape appears comparatively conserved and may be less informative than fruit shape or seed-coat colour in distinguishing superior material. The uneven distribution of fruit shapes among populations has clear conservation implications. Some shapes were widespread, whereas others were rare and locally restricted, indicating that baobab diversity in Malawi is geographically distributed rather than concentrated in a single site. Conservation strategies focused on only one or two populations would therefore risk missing part of the country’s phenotypic diversity. This supports maintaining multiple populations in situ while also sampling broadly for ex situ conservation and domestication. For agroforestry development, that is an important point, improvement efforts should not focus only on the highest-yielding sites, but should also capture rare and potentially valuable phenotypes from other locations. Because fruit shapes were sampled in a balanced way, the diversity metrics reported here should be interpreted as indicators of trait representation rather than natural abundance. The PCA reinforces the view that baobab variation is structured along more than one functional axis. One axis reflected overall fruit productivity, whereas another captured individual seed morphology. This indicates that selecting for high fruit weight will not necessarily improve all other desirable traits. Similar multivariate structuring has been reported in West African baobab populations (Assogbadjo et al. 2006 ; De Smedt et al. 2011 ). For domestication and agroforestry management, the implication is straightforward, selection criteria should be matched to the intended end use from the outset. Fruit weight may be a useful field proxy where pulp production is the priority, but seed-based value chains will require additional screening for seed traits. Overall, the study supports a targeted rather than generic approach to baobab domestication in Malawi. Wild and parkland populations contain substantial phenotypic diversity, but that diversity is structured in ways that matter for use, management, and conservation. High spheroid and oblong fruits appear promising for pulp-oriented improvement, whereas phenotypes with larger individual seeds may be more suitable for seed-based products. For agroforestry development, the main lesson is that domestication goals should be defined clearly before selection begins. Future work should test the stability of these traits under common garden or clonal trials and assess whether seed-coat colour is associated with germination or biochemical differences. Integrating such phenotypic screening with emerging genomic tools would provide a stronger basis for baobab improvement and conservation in smallholder agroforestry landscapes (Kitony et al. 2024 ). 5.0 Conclusion Wild and parkland baobab populations in Malawi harbor substantial and structured phenotypic variation in fruit and seed traits, making them valuable for domestication, germplasm development, and conservation. The diversity of fruit shapes, including previously unreported forms, highlights the importance of these populations as reservoirs of under characterized variation. Because fruit shape is linked to productivity, and seed traits vary in ways relevant to different end uses, baobab improvement should rely on targeted multi-trait selection rather than a single productivity measure. The geographically distributed nature of this diversity also supports multi-site conservation and broad germplasm sampling. Together, these findings provide a practical basis for baobab domestication in Malawi and reinforce the need to conserve wild populations while advancing improvement through common garden testing and genomic tools. Declarations Acknowledgements This work was supported by the Regional Universities Forum for Capacity Building in Agriculture (RUFORUM). The authors are indebted to RUFORUM for funding this project (Grant Number: RU 2014 GRG 107), and the views expressed here are not necessarily those of RUFORUM. Above all, we would like to pay our gratitude and our respects to our late colleague Prof. Dr. Chimuleke R. Y. Munthali. After helping this research Prof. Munthali sadly passed away before we could think of publishing this work. He was a dedicated professor in the Department of Forestry & Environmental Management at Mzuzu University, Malawi, with a passion for research on the baobab tree. Declaration of Competing Interests The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. Author Contributions Herbert Jenya: conceptualization, methodology, investigation, data curation, analysis, writing original draft, writing review and editing. Owen Kachala: analysis, writing review and editing. Data Availability Statement The data that support the findings of this study are available from the corresponding author (Herbert Jenya; [email protected] ) upon reasonable request. Data include fruit and seed morphological measurements, tree diameter at breast height records, and tree morphological trait observations collected across five baobab populations in Malawi. References Abere M, Yilma Z, Tsegie T, Eshete A, Alemu A (2023) Population structure and phenological attributes of Adansonia digitata L. (baobab) in Northwestern lowland area of Ethiopia. Heliyon 9:e22571. https://doi.org/10.1016/j.heliyon.2023.e22571 Anjarwalla P, Ofori D, Owino A, Matuku D, Adika W, Njogu K, Kehlenbeck K (2017) Testing different grafting methods for vegetative propagation of baobab (Adansonia digitata L.) in Kenya to assist its domestication and promote cultivation. Forests Trees and Livelihoods 26:85–95. https://doi.org/10.1080/14728028.2016.1202791 Assogbadjo AE, Chadare FJ, Manda L, Sinsin B (2021) A 20-year journey through an orphan African baobab (Adansonia digitata L.) towards improved food and nutrition security in Africa. Frontiers in Sustainable Food Systems 5:675382. https://doi.org/10.3389/fsufs.2021.675382 Assogbadjo AE, Glèlè Kakaï R, Edon S, Kyndt T, Sinsin B (2011) Natural variation in fruit characteristics, seed germination and seedling growth of Adansonia digitata L. in Benin. New Forests 41:113–125. https://doi.org/10.1007/s11056-010-9214-z Assogbadjo AE, Kyndt T, Sinsin B, Gheysen G, Van Damme P (2006) Patterns of genetic and morphometric diversity in baobab (Adansonia digitata): populations across different climatic zones of Benin (West Africa). Annals of Botany 97:819–830. https://doi.org/10.1093/aob/mcl043 Assogbadjo AE, Sinsin B, Codjia JTC, Van Damme P (2005) Ecological diversity and pulp, seed and kernel production of the baobab (Adansonia digitata) in Benin. Belgian Journal of Botany 138:47–56 Assogba D, Idohou R, Chirwa P, Assogbadjo AE (2022) On opportunities and challenges to conserve the African baobab under present and future climates in Benin (West Africa). Journal of Arid Environments 198:104692. https://doi.org/10.1016/j.jaridenv.2021.104692 Buchmann C, Prehsler S, Hartl A, Vogl CR (2010) The importance of baobab (Adansonia digitata L.) in rural West African subsistence: suggestion of a cautionary approach to international market export of baobab fruits. Ecology of Food and Nutrition 49:145–172. https://doi.org/10.1080/03670241003766014 Chládová A, Kalousová M, Mandák B, Kehlenbeck K, Prinz K, Šmíd J, Van Damme P, Lojka B (2019) Genetic diversity and structure of baobab (Adansonia digitata L.) in southeastern Kenya. Royal Society Open Science 6:190854. https://doi.org/10.1098/rsos.190854 Cossam FG, Dzanja JK, Kamwana B, Olumeh DE, Mithöfer D (2023) Determinants of postharvest losses along the baobab value chain in Malawi. Frontiers in Sustainable Food Systems 7:1119107. https://doi.org/10.3389/fsufs.2023.1119107 Cuni Sanchez A, De Smedt S, Haq N, Samson R (2011) Comparative study on baobab fruit morphological variation between western and south-eastern Africa: opportunities for domestication. Genetic Resources and Crop Evolution 58:1143–1156. https://doi.org/10.1007/s10722-010-9647-4 Cuni Sanchez A, Haq N, Assogbadjo AE (2010) Variation in baobab leaf morphology and its relation to drought tolerance. Genetic Resources and Crop Evolution 57:17–25. https://doi.org/10.1007/s10722-009-9447-x de Mendiburu F (2023) agricolae: statistical procedures for agricultural research. R package version 1.3-7. https://doi.org/10.32614/CRAN.package.agricolae De Smedt S, Alaerts K, Kouyate AM, Van Damme P, Potters G, Samson R (2011) Phenotypic variation of baobab (Adansonia digitata L.) fruit traits in Mali. Agroforestry Systems 82:87–97. https://doi.org/10.1007/s10457-010-9357-0 Dixon RA, Sumner LW (2003) Legume natural products: understanding and manipulating complex pathways for human and animal health. Plant Physiology 131:878–885. https://doi.org/10.1104/pp.102.017319 Gebauer J, Luedeling E (2013) A note on baobab (Adansonia digitata L.) in Kordofan, Sudan. Genetic Resources and Crop Evolution 60:1587–1596. https://doi.org/10.1007/s10722-013-9964-5 Gruenwald J (2009) Novel botanical ingredients for beverages. Clinics in Dermatology 27:210–216 Government of the Republic of Malawi (2005) Food security and nutrition project: project appraisal report. Food and Agriculture Organization of the United Nations Gruenwald J, Galizia M (2005) Adansonia digitata: market brief in the European Union for selected natural ingredients derived from native species. United Nations Conference on Trade and Development Gurashi NA, Kordofani MA (2014) Morphological variation in fruit shapes of Adansonia digitata L. from Blue Nile and North Kordofan States. Journal of Forest Products and Industries 3:106–111 Jenya H, Munthali CRY, Mhango J (2018) Amenability of African baobab (Adansonia digitata L.) to vegetative propagation techniques. Journal of Sustainable Forestry. https://doi.org/10.1080/10549811.2018.1449123 Kalinganire A, Savadogo P, Ky-Dembele C, Bush D, Ræbild A, Tougiani A, Muchugi A, Traore FT, Kone B (2023) Evidence of early genetic variation in survival and growth traits of baobab (Adansonia digitata L.): the way forward for domestication and breeding. Agroforestry Systems 97:1221–1231. https://doi.org/10.1007/s10457-023-00837-z Kamatou GPP, Vermaak I, Viljoen AM (2011) An updated review of Adansonia digitata: a commercially important African tree. South African Journal of Botany 77:908–919. https://doi.org/10.1016/j.sajb.2011.08.010 Kehlenbeck K, Padulosi S, Alercia A (2015) Descriptors of baobab (Adansonia digitata L.). Bioversity International and World Agroforestry Centre Kitony JK, Colt K, Abramson BW, Hartwick NT, Petrus S, Konozy EHE, Karimi N, Yant L, Michael TP (2024) Chromosome-level baobab genome illuminates its evolutionary trajectory and environmental adaptation. Nature Communications 15:8833. https://doi.org/10.1038/s41467-024-53157-w Lê S, Josse J, Husson F (2008) FactoMineR: an R package for multivariate analysis. Journal of Statistical Software 25:1–18. https://doi.org/10.18637/jss.v025.i01 Luckert KM, Nemarundwe N, Gibbs L, Grundy I, Hauer D, Maruzane D, Shackleton S, Sithole J (2014) Contribution of baobab production activities to household livelihoods: a case study in Zimbabwe. In: Cunningham BA, Campbell MB, Luckert KM (eds) Bark use, management, and commerce in Africa. Advances in Economic Botany, vol 17. New York Botanical Garden Press, pp 14 Meinhold K, Darr D (2022) Keeping up with rising (quality) demands? The transition of a wild food resource to mass market, using the example of baobab in Malawi. Frontiers in Sustainable Food Systems 6:840760. https://doi.org/10.3389/fsufs.2022.840760 Munthali CRY (2012) Use, physiology and genetic characterisation of selected natural populations of Adansonia digitata in Malawi. Doctoral dissertation, Stellenbosch University Munthali CRY, Chirwa PW, Akinnifesi FK (2012) Phenotypic variation in fruit and seed morphology of Adansonia digitata L. (baobab) in five selected wild populations in Malawi. Agroforestry Systems 85:279–290. https://doi.org/10.1007/s10457-012-9500-1 Ngole MA, Lusambo LP, Temu BJ (2024) Baobab trees population and economic contribution: stems density, asset value and fruits value chains in Kilolo and Iringa Urban districts, Tanzania. Trees, Forests and People 16:100519. https://doi.org/10.1016/j.tfp.2024.100519 Ochuodho JO, Modi AT (2008) Dormancy of wild mustard (Sisymbrium capense) seeds is related to seed coat colour. Seed Science and Technology 36:46–55. https://doi.org/10.15258/sst.2008.36.1.05 Oksanen J, Simpson G, Blanchet F, Kindt R, Legendre P, Minchin P, O’Hara R, Solymos P, Stevens M, Szoecs E, Wagner H et al (2026) vegan: community ecology package. R package version 2.7-3. https://doi.org/10.32614/CRAN.package.vegan Omondi M, Rimberia FK, Wainaina CM, Mukundi JBN, Orina J, Gebauer J, Kehlenbeck K (2019) Fruit morphological diversity and productivity of baobab (Adansonia digitata L.) in coastal and lower eastern Kenya. Forests Trees and Livelihoods 28:266–280. https://doi.org/10.1080/14728028.2019.1659861 Patrut A, Woodborne S, Patrut RT, Rakosy L, Lowy DA, Hall G, Von Reden KF (2018) The demise of the largest and oldest African baobabs. Nature Plants 4:423–426 R Core Team (2025) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna Rashidi M, Seyif K (2007) Classification of fruit shape in cantaloupe using analysis of geometrical attributes. World Journal of Agricultural Sciences 3:735–740 Santos RC, Pires JL, Correa RX (2012) Morphological characterization of leaf, flower, fruit, and seed traits among Brazilian Theobroma L. species. Genetic Resources and Crop Evolution 59:327–345 Sidibé M, Williams JT (2002) Baobab: Adansonia digitata L. International Centre for Underutilised Crops, Southampton Stadlmayr B, Wanangwa J, Waruhiu C, Jamnadass R, Kehlenbeck K (2020) Nutritional composition of baobab (Adansonia digitata L.) fruit pulp sampled at different geographical locations in Kenya. Journal of Food Composition and Analysis 94:103617. https://doi.org/10.1016/j.jfca.2020.103617 Tchoundjeu Z, Atangana A, Asaah E, Tsobeng A, Facheux C, Foundjem D, Mbosso C, Degrande A, Sado T, Kanmegne J, Mbile P, Tabuna H, Anegbeh P, Useni M (2008) Domestication, utilization and marketing of indigenous fruit trees in West and Central Africa. In: Akinnifesi FK, Leakey RRB, Ajayi OC, Sileshi G, Tchoundjeu Z, Matakala P, Kwesiga FR (eds) Indigenous fruit trees in the tropics: domestication, utilization and commercialization. CAB International, pp 137–170 Tremayne MA, Richards AJ (2000) Seed weight and seed number affect subsequent fitness in outcrossing and selfing in Primula species. New Phytologist 148:127–142 Venter SM, Witkowski ETF (2010) Baobab (Adansonia digitata L.) density, size-class distribution and population trends between four land-use types in northern Venda, South Africa. Forest Ecology and Management 259:294–300. https://doi.org/10.1016/j.foreco.2009.10.016 Wickens GE, Lowe P (2008) The baobabs: the pachycauls of Africa, Madagascar and Australia. Springer Additional Declarations No competing interests reported. Cite Share Download PDF Status: Under Review Version 1 posted Editorial decision: Revision requested 29 Apr, 2026 Reviews received at journal 24 Apr, 2026 Reviews received at journal 23 Apr, 2026 Reviewers agreed at journal 14 Apr, 2026 Reviewers agreed at journal 13 Apr, 2026 Reviews received at journal 12 Apr, 2026 Reviewers agreed at journal 11 Apr, 2026 Reviewers invited by journal 10 Apr, 2026 Editor assigned by journal 07 Apr, 2026 Submission checks completed at journal 04 Apr, 2026 First submitted to journal 03 Apr, 2026 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-9317216","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":623399878,"identity":"e02a764b-d4d8-4656-992d-ae30c8d6a899","order_by":0,"name":"Herbert Jenya","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA/0lEQVRIiWNgGAWjYBACAwYGNhDNDOZ9AGI2duK0GIC1MM4AaWEmUguYw8yDsBA3MGfvffbg454/7PzSPYafbX5tk+djZmD88DEHtxbLnuPmhjOeGTBLzjljLJ3bd9uwjZmBWXLmNjwOu5HGJs1zwIDZ4EaOgXRuz21GoBY2Zl58Wu4/Y5P+A9RifyPH+Ldlz217wlpusLFJM4Bskcgxk2b4cTuRsJYzaeyGPQeMmSVupJVZ9jbcTm5jZmzG75fjx9ge/Dggl8w/I3nzjR9/btvOb28++OEjHi0wkMzAwGHAwNgGYjM2EFYPBHYMDOwPGBj+EKV4FIyCUTAKRhgAANMrTOLnF1oSAAAAAElFTkSuQmCC","orcid":"","institution":"Forestry Research Institute of Malawi","correspondingAuthor":true,"prefix":"","firstName":"Herbert","middleName":"","lastName":"Jenya","suffix":""},{"id":623399879,"identity":"6a610a4b-4afc-418a-9db5-ab4116b5737c","order_by":1,"name":"Owen Kachala","email":"","orcid":"","institution":"Forestry Research Institute of Malawi","correspondingAuthor":false,"prefix":"","firstName":"Owen","middleName":"","lastName":"Kachala","suffix":""}],"badges":[],"createdAt":"2026-04-04 03:38:30","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-9317216/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9317216/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":107180537,"identity":"93d3ecfd-cb85-4e03-b783-e9e5a1c3555d","added_by":"auto","created_at":"2026-04-17 16:59:53","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":405259,"visible":true,"origin":"","legend":"\u003cp\u003eDiversity of fruit shapes in five baobab populations in Malawi. (1=Ellipsoid, 2=Oblong, 3=Ovate, 4=Obovate, 5=Fusiform, 6=High spheroid, 7=Globose, 8=Ellipsoid pointed, 9=Oblong compressed, 10=Oblong pointed, 11=Rhomboid, 12=Spheroid emarginate, 13=Clavate)\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-9317216/v1/8249471a77b357664aaae46a.png"},{"id":107483302,"identity":"1709a91c-40e4-4d6d-84bc-44d6c6252666","added_by":"auto","created_at":"2026-04-22 02:27:16","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":189133,"visible":true,"origin":"","legend":"\u003cp\u003eSeed colours in five baobab populations in Malawi (A=Light brown, B=Brown, C=Dark brown and D=Greyish brown)\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-9317216/v1/1e5efbcc7a056bd8c9d7fed3.png"},{"id":107180539,"identity":"8c436099-4d8e-4e7a-b20b-69910286b2fe","added_by":"auto","created_at":"2026-04-17 16:59:53","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":82303,"visible":true,"origin":"","legend":"\u003cp\u003eSeed shape in five baobab populations in Malawi (A=Very-reniform, B=Reniform and C= Oblong)\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-9317216/v1/872beaab037bcef8217be591.png"},{"id":107180540,"identity":"cff999f9-d6d6-41ca-b801-fb438a4bd31b","added_by":"auto","created_at":"2026-04-17 16:59:53","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":320188,"visible":true,"origin":"","legend":"\u003cp\u003ePrincipal Component Analysis (PCA) of 11 quantitative fruit and seed traits of baobab (Adansonia digitata L.) across five populations in Malawi. (A) Biplot showing individual fruit scores coloured and shaped by population; dashed ellipses represent 95% confidence regions; crosses (+) mark population centroids; arrows indicate variable loadings.\u003c/p\u003e","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-9317216/v1/674ba4f5e96fec527d680d45.png"},{"id":107482596,"identity":"9fb8a764-650e-452f-b016-a02a0038dc87","added_by":"auto","created_at":"2026-04-22 02:24:09","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":179754,"visible":true,"origin":"","legend":"\u003cp\u003eTree morphological characteristics of baobab (Adansonia digitata L.) across five districts in Malawi: (A) bark colour, (B) tree shape, (C) trunk shape, and (D) growth habit.\u003c/p\u003e","description":"","filename":"floatimage5.png","url":"https://assets-eu.researchsquare.com/files/rs-9317216/v1/b359678c3de73e103992c3e9.png"},{"id":107486027,"identity":"d0a8eba3-d1ba-4879-8c06-c4094981a175","added_by":"auto","created_at":"2026-04-22 02:37:14","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2125519,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9317216/v1/fda8ba92-60dd-4391-8c57-3ba7c2e24663.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Trait diversity in wild and parkland baobab (Adansonia digitata L.) populations in Malawi","fulltext":[{"header":"1.0 INTRODUCTION","content":"\u003cp\u003e \u003cem\u003eAdansonia digitata\u003c/em\u003e L. belongs to the subfamily Bombacoideae within the family Malvaceae. The genus \u003cem\u003eAdansonia\u003c/em\u003e comprises eight species, of which \u003cem\u003eA. digitata\u003c/em\u003e is the widespread African species (Wickens \u0026amp; Lowe, \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Commonly known as baobab, monkey bread tree, or bottle tree, it is one of Africa\u0026rsquo;s most iconic long-lived tree species and is valued for its ecological, cultural, and economic importance (Kehlenbeck et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Sidib\u0026eacute; and Williams \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Wickens and Lowe \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). The tree is characterized by massive trunks, broad spreading branches, and a large rounded canopy (Sidib\u0026eacute; and Williams \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). Some individuals are exceptionally old, with ages extending well beyond a millennium (Patrut et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOver the past two decades, increasing research attention has positioned baobab as an important orphan tree species for food and nutrition security, livelihood support, and agroforestry development (Assogbadjo et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In Malawi, baobab occurs mainly in wild stands in savannah landscapes and on farmland. Baobab remains important in local diets and as a source of income for rural communities (Gebauer and Luedeling \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Kamatou et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Numerous products are made locally, including baobab pulp, sweets, ice lollies, juice, jam, coffee substitutes, soap, and lip balm (Munthali \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Additionally, products in wider markets include beverages, cereals, cereal bars, probiotic yogurts, and cosmetic products derived from baobab seed oil (Gruenwald \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Gruenwald and Galizia \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Kamatou et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Different plant parts, including the roots, bark, leaves, and seeds, are also used in traditional medicine to treat various ailments (Buchmann et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Wickens and Lowe \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Recent studies further show that baobab fruit pulp is rich in key nutrients and that its composition varies geographically, reinforcing the species\u0026rsquo; growing importance in food and nutrition research as well as the need to identify superior germplasm (Assogbadjo et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Stadlmayr et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe commercialization of baobab products continues to contribute to rural livelihoods in areas where the species is abundant. Recent studies show that its value now extends beyond household use to increasingly organized local and regional value chains. In Malawi, the baobab sector has in recent years shifted from mainly subsistence use toward wider business opportunity, although this transition remains constrained by weak policy support, rising quality requirements, and uneven value chain coordination (Meinhold et al. 2022). Recent evidence from Tanzania likewise shows that baobab fruit value chains generate economic value through margins, wages, and taxes, but are limited by poor storage, low prices, lack of capital, poor processing facilities, seasonal fruit supply, and limited access to market information (Ngole et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Studies from Malawi further show that postharvest losses reduce the value ultimately captured by collectors, traders, and processors, highlighting inefficiencies in the chain (Cossam et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). These newer findings complement earlier evidence that baobab trade contributes meaningfully to household livelihoods in southern Africa (Luckert et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). This implies that the long-term economic potential of baobab will depend on stronger value chain organization, reduced losses, better processing and storage, and more deliberate domestication and management of the resource base.\u003c/p\u003e \u003cp\u003eConservation concerns have become increasingly important in recent years. Emerging evidence suggests that baobab populations in several parts of Africa are under pressure from land-use change, agricultural expansion, climate variability, and weak natural regeneration (Abere et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Assogba et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Venter and Witkowski \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Studies on habitat suitability under present and future climates indicate that persistence of the species will depend not only on protecting existing stands, but also on supporting regeneration, restoration, and cultivation within farming systems (Assogba et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). In addition, recent syntheses emphasize that, despite growing interest in baobab, the species remains underexploited relative to its potential and still faces important biological and management constraints that could limit its contribution to food systems and rural livelihoods if conservation and domestication efforts are not strengthened (Assogbadjo et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eKnowledge of trait variation in baobab is crucial for selecting plant material with desirable characteristics for domestication, conservation, and improvement (Santos et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Previous studies have documented phenotypic variation in baobab fruits, leaves, seeds, and tree traits in Malawi and elsewhere in Africa (Assogbadjo et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2005\u003c/span\u003e, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2006\u003c/span\u003e, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Cuni Sanchez et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2010\u003c/span\u003e, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; De Smedt et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Munthali et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Omondi et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). More recent work also shows substantial genetic diversity and population structure in \u003cem\u003eA. digitata\u003c/em\u003e, strengthening the case for conserving diverse germplasm while identifying superior material for domestication (Chl\u0026aacute;dov\u0026aacute; et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Likewise, evidence of early genetic variation in survival and growth traits indicates that baobab improvement and breeding are feasible and can support future domestication efforts (Kalinganire et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Fruit shape may be an especially important trait because it may influence consumer preference, marketability, productivity, and the development of ideotypes for domestication (Rashidi and Seyfi 2007; Vihotogb\u0026eacute; et al. 2013). Multivariate approaches such as principal component analysis and cluster analysis have proven useful in identifying independent axes of phenotypic variation and grouping baobab populations for domestication prioritisation (Assogbadjo et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; De Smedt et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Tree morphological characteristics, including diameter at breast height (DBH), also vary among populations and may reflect differences in population age structure, management history, and land-use context (Assogbadjo et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Venter and Witkowski \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2010\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eHowever, important gaps remain for Malawi. Although previous studies have examined fruit and seed traits in selected wild populations (Munthali et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), information is still limited on the diversity of fruit shapes across populations, the relationship between fruit shape and quantitative fruit and seed traits, and the extent to which fruit productivity and seed morphology represent independent axes of variation. An integrated characterisation that combines fruit shape diversity indices, population-level trait variation, multivariate structure, and tree morphological attributes is therefore needed to support conservation, domestication, and selection of superior germplasm in Malawian baobab populations.\u003c/p\u003e \u003cp\u003eThis study aimed to characterise the trait diversity of baobab across five wild and parkland populations in Malawi to inform conservation, management, improvement, and domestication efforts. Specifically, it addressed the following questions: (1) What fruit shapes occur, and do they differ consistently between individual trees and among populations? (2) Do quantitative fruit traits vary among fruit shapes? (3) Do quantitative and qualitative seed traits vary among fruit shapes? (4) How does fruit shape diversity, quantified using the Shannon\u0026ndash;Wiener diversity index, compare across the five populations? (5) Do fruit and seed traits differ significantly among populations? (6) What is the multivariate structure of fruit and seed phenotypic variation, and are fruit productivity and seed morphology independent axes of variation? and (7) How do tree morphological characteristics and diameter at breast height vary among populations? We hypothesize that baobab populations in Malawi exhibit significant phenotypic variation in fruit shape, fruit traits, seed traits, and tree morphology. Specifically, fruit-shape categories are expected to differ in associated quantitative fruit and seed characteristics, populations are expected to differ in fruit-shape diversity and overall trait composition, and multivariate analysis is expected to reveal distinct dimensions of variation separating fruit productivity traits from seed morphology traits.\u003c/p\u003e"},{"header":"2.0 MATERIALS AND METHODS","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1 Study site\u003c/h2\u003e \u003cp\u003eFruit samples were collected from five baobab populations in Chikwawa, Karonga, Mangochi, Neno, and Salima (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Study populations were selected to represent the major agro-ecological settings across Malawi in which the species occurs, based on national agro-ecological classifications reported by Government of Malawi (Government of the Republic of Malawi \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). The sites span considerable variation in altitude, mean annual rainfall (MAR), temperature, and soil type (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Chikwawa, situated in the Lower Shire Valley, is characterised by low elevation (\u0026lt;\u0026thinsp;200 m), MAR of 710\u0026ndash;850 mm, mean annual temperatures exceeding 25\u0026deg;C, and predominantly vertisol soils. Mangochi and Salima, both located in the lakeshore plains, receive similar rainfall (710\u0026ndash;850 mm) with alluvial calcimorphic soils overlying vertisols and mean annual temperatures of 20\u0026ndash;25\u0026deg;C. Neno, in the southern highlands, receives 1,200\u0026ndash;1,600 mm of rainfall annually with sandy ferralitic soils and cooler temperatures of 19\u0026ndash;21\u0026deg;C. Karonga, in the northern lakeshore zone, is the wettest site (MAR\u0026thinsp;\u0026gt;\u0026thinsp;1,600 mm) with predominantly ferrosols and regosols and mean annual temperatures of 23\u0026ndash;25\u0026deg;C (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Further, populations were selected based on abundance of trees and prevailing trade in baobab products.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eLocation, physical description of baobab populations and number of trees sampled in Malawi\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePopulation\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eCoordinates\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAgro-ecological zone\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAltitude\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eMean annual rainfall (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eMean annual temperature\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSoil type\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eChikwawa (21**)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e16\u0026deg;02\u0026rsquo;S, 34\u0026deg;50\u0026rsquo;E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eShire Valley\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;200\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e710\u0026ndash;850\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u0026gt;\u0026thinsp;25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eVertisols\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMangochi (30**)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14\u0026deg;25\u0026rsquo;S, 35\u0026deg;16\u0026rsquo;E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLakeshore plains\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e200\u0026ndash;1200\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e710\u0026ndash;850\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e20\u0026ndash;25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eAlluvial calcimorphic soils above the vertisols\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSalima (30**)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e13\u0026deg;47\u0026rsquo;S, 34\u0026deg;28\u0026rsquo;E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLakeshore plains\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e200\u0026ndash;1200\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e710\u0026ndash;850\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e20\u0026ndash;25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eAlluvial calcimorphic soils above the vertisols\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNeno (30**)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e15\u0026deg;25\u0026rsquo;S, 34\u0026deg;40\u0026rsquo;E\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSouthern Highlands\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e900\u0026ndash;1500\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1200\u0026ndash;1600\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e19\u0026ndash;21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSandy ferralitic\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKaronga (24**)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e09\u0026deg;56ˈS 33\u003csup\u003e\u0026deg;\u003c/sup\u003e56ˈE\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNorthern Lakeshore\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e475\u0026ndash;1000\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u0026gt;\u0026thinsp;1600\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e23\u0026ndash;25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eFerrosols dominant; regosols\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"7\"\u003e** number of trees sampled\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.2 Data collection\u003c/h2\u003e \u003cp\u003eFruits were collected at the peak of fruit maturity between March and May of 2016. Dry fruits (10 per tree) were randomly picked from trees. A total of 135 trees were characterized from the five populations (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFruit shape description was done according to Gurashi and Kordofani (\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) and Kehlenbeck et al. (\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). From each fruit shape, subsamples of five fruits were randomly selected for the assessment of fruit, seed, and pulp weight. Fruits were weighed to the nearest 1 g with an electronic scale (TREE LW Measurements HRB 10001-High Resolution Balance 10,000 g \u0026times;0.1 g). Fruit shells were cracked manually and the fruit content (pulp\u0026thinsp;+\u0026thinsp;seeds) were removed and weighed moist. Pulp was separated from seeds by washing with water, and seeds were then air dried and weighed again. Pulp content was calculated by subtracting the moist seed weight from the fruit content (pulp\u0026thinsp;+\u0026thinsp;seeds). Seeds were counted to determine the seed number per fruit. Single seed weight was calculated by dividing the seed weight by the number of seeds according to Cuni Sanchez et al., (\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). A subsample of ten seeds per fruit was randomly selected for the measurement of seed length, seed width, and seed thickness using a micro-calliper to the nearest mm. Seed shape and colour were visually assessed in the laboratory using the guide proposed by Kehlenbeck et al. (\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2015\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTree morphological traits were recorded for each of the 135 sampled trees at the time of fruit collection. Diameter at breast height (DBH) was measured at 1.3 m above ground level using a diameter tape. Crown shape, trunk shape, bark colour, and bark texture were assessed visually for each tree following the descriptors proposed by Kehlenbeck et al. (\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Crown shape was classified as roundish, ellipsoid, or semi-circular. Trunk shape was categorised as cylindrical, bell-shaped, concave, or cone-shaped. Bark colour was recorded as grey or reddish-brown, and bark texture as smooth or rough. The branch growth habit of each tree was recorded as erect, spreading, or drooping. The land use context of each tree was also noted, classifying trees as occurring in parkland, fallow land, forest/woodland, backyard garden, or shrubland.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e2.3 Statistical analysis\u003c/h2\u003e \u003cp\u003eFruit shape occurrence was analysed using descriptive statistics. Quantitative fruit and seed traits were compared among fruit shapes and among populations (districts) using one-way ANOVA with Sidak pairwise comparisons and compact letter displays were generated using the agricolae package (de Mendiburu \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) in R (R Core Team 2024). Principal Component Analysis (PCA) was performed on standardised quantitative traits to examine the multivariate structure of variation among populations, using the FactoMineR package (L\u0026ecirc; et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Fruit shape diversity within each population was quantified using the Shannon-Wiener diversity index (H' = \u0026minus;Σ p\u003csub\u003ei\u003c/sub\u003e ln p\u003csub\u003ei\u003c/sub\u003e), where p\u003csub\u003ei\u003c/sub\u003e is the relative frequency of each fruit shape, and Pielou's evenness (J' = H'/ln S, S\u0026thinsp;=\u0026thinsp;number of shapes), computed using the vegan package (Oksanen et al. 2024). Tree morphological traits were summarised using frequency distributions by district. Diameter at breast height (DBH) was compared among districts using one-way ANOVA with Sidak pairwise comparisons. All statistical analyses were conducted in R version 4.5.2 (R Core Team 2024).\u003c/p\u003e \u003c/div\u003e"},{"header":"3.0 RESULTS","content":"\u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003e3.1 Fruit Shape Diversity among Populations\u003c/h2\u003e \u003cp\u003eThe Shannon\u0026ndash;Wiener diversity index was highest in Chikwawa and Neno, which supported the highest fruit shape richness and diversity (H' = 2.30, S\u0026thinsp;=\u0026thinsp;10 shapes each), followed by Karonga (H' = 2.08, S\u0026thinsp;=\u0026thinsp;8). Mangochi and Salima had the fewest shapes (H' = 1.95, S\u0026thinsp;=\u0026thinsp;7). Pielou's evenness was J' = 1.00 in all populations, a reflection of the balanced sampling design rather than a biological pattern (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eFruit shape diversity indices for five baobab populations in Malawi.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePopulation\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eShannon (H')\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNo. of shapes (S)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eEvenness (J')\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eChikwawa\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2.30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKaronga\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2.08\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMangochi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1.95\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNeno\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e2.30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSalima\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1.95\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"4\"\u003eNote: Evenness (J' = 1) reflects balanced sampling design\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eThis study identified thirteen distinct fruit shapes across the sampled baobab populations, namely ellipsoid, oblong, ovate, fusiform, high spheroid, globose, ellipsoid pointed, oblong compressed, oblong pointed, rhomboid, spheroid emarginate, and clavate (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e; Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Ellipsoid, oblong, and ovate were common to all five populations, while spheroid emarginate was unique to Mangochi (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eFruit shape occurrence (%) in five baobab populations in Malawi\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eFruit shape\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eChikwawa (A)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eKaronga (L)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMangochi (Ba)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNeno (J)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSalima (Ba)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSouth\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNorth\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSouth\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSouth\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eCentral\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eClavate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEllipsoid\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e43\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEllipsoid pointed\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFusiform\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eHigh spheroid\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026mdash;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eOblong\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e24\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eOblong compressed\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eOblong pointed\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eObovate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eOvate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRhomboid\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSpheroid\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSpheroid emarginate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u0026mdash;0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003e3.2 Variation of Fruit and Seed Traits among Fruit Shapes\u003c/h2\u003e \u003cp\u003eEleven fruit and seed traits measured differed significantly (\u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026le;\u0026thinsp;0.001) among the 13 fruit shapes (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). High spheroid fruits were consistently the largest by weight-related measures heaviest (235\u0026thinsp;\u0026plusmn;\u0026thinsp;17 g), widest (9.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3 cm), highest pulp content (45.3\u0026thinsp;\u0026plusmn;\u0026thinsp;3.9 g), greatest seed mass (98.1\u0026thinsp;\u0026plusmn;\u0026thinsp;8.1 g), and most seeds per fruit (208\u0026thinsp;\u0026plusmn;\u0026thinsp;16) differing significantly from fusiform and ovate fruits in most traits. Oblong fruits ranked second for most weight metrics. In contrast, fusiform (119\u0026thinsp;\u0026plusmn;\u0026thinsp;17 g) and ovate (131\u0026thinsp;\u0026plusmn;\u0026thinsp;15 g) fruits were the lightest, narrowest, and had the fewest seeds. Fruit length followed a different pattern: oblong pointed (19.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9 cm), ellipsoid pointed (18.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6 cm), and clavate (17.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9 cm) were the longest, while spheroid (10.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.7 cm) and spheroid emarginate (10.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.3 cm) were the shortest. At the individual seed level, clavate fruits consistently produced the largest seeds across all four dimensions heaviest single seeds (0.59\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 g), longest (12.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2 mm), widest (9.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2 mm), and thickest (7.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1 mm) while ovate and spheroid fruits consistently had the smallest seeds. Seed dimensions showed notably less variation among shapes than fruit-level traits (CVs 6\u0026ndash;22% vs. 19\u0026ndash;52%). Full means and Sidak groupings for all eleven traits are presented in Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eVariation of fruit and seed traits (mean\u0026thinsp;\u0026plusmn;\u0026thinsp;SE) among the 13 fruit shapes in five baobab populations in Malawi. Means in the same column sharing the same letter do not differ significantly (Sidak, p\u0026thinsp;\u0026lt;\u0026thinsp;0.05). FW\u0026thinsp;=\u0026thinsp;fruit weight, FL\u0026thinsp;=\u0026thinsp;fruit length, FWd\u0026thinsp;=\u0026thinsp;fruit width, PW\u0026thinsp;=\u0026thinsp;pulp weight, % Pulp = % pulp to fruit weight, TSW\u0026thinsp;=\u0026thinsp;total seed weight, NS\u0026thinsp;=\u0026thinsp;number of seeds, SSW\u0026thinsp;=\u0026thinsp;single seed weight, SL\u0026thinsp;=\u0026thinsp;seed length, SWd\u0026thinsp;=\u0026thinsp;seed width, ST\u0026thinsp;=\u0026thinsp;seed thickness.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"12\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eFruit shape\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"5\" nameend=\"c6\" namest=\"c2\"\u003e \u003cp\u003eFruit traits\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"6\" nameend=\"c12\" namest=\"c7\"\u003e \u003cp\u003eSeed traits\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eFW (g)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFL (cm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFWd (cm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003ePW (g)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003e% Pulp\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eTSW (g)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eNS\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eSSW (g)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003eSL (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c11\"\u003e \u003cp\u003eSWd (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c12\"\u003e \u003cp\u003eST (mm)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eClavate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e168.4\u0026thinsp;\u0026plusmn;\u0026thinsp;20.9abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e17.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e27.0\u0026thinsp;\u0026plusmn;\u0026thinsp;4.8ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e15.52\u0026thinsp;\u0026plusmn;\u0026thinsp;1.53a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e59.3\u0026thinsp;\u0026plusmn;\u0026thinsp;10.0abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e99\u0026thinsp;\u0026plusmn;\u0026thinsp;19ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.59\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e12.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e9.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e7.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEllipsoid\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e182.4\u0026thinsp;\u0026plusmn;\u0026thinsp;13.2abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e16.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6cd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e8.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e33.0\u0026thinsp;\u0026plusmn;\u0026thinsp;3.0ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e17.90\u0026thinsp;\u0026plusmn;\u0026thinsp;0.97a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e68.6\u0026thinsp;\u0026plusmn;\u0026thinsp;6.3abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e142\u0026thinsp;\u0026plusmn;\u0026thinsp;12abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.50\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e11.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1abcd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e8.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e7.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1ab\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEllipsoid pointed\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e187.4\u0026thinsp;\u0026plusmn;\u0026thinsp;13.2abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e18.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e8.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2cd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e32.0\u0026thinsp;\u0026plusmn;\u0026thinsp;3.0ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e17.07\u0026thinsp;\u0026plusmn;\u0026thinsp;0.97a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e69.4\u0026thinsp;\u0026plusmn;\u0026thinsp;6.3abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e129\u0026thinsp;\u0026plusmn;\u0026thinsp;12ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.54\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e12.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1cd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e9.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e7.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1ab\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eFusiform\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e118.7\u0026thinsp;\u0026plusmn;\u0026thinsp;17.1a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e17.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.7d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e24.7\u0026thinsp;\u0026plusmn;\u0026thinsp;3.9a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e19.34\u0026thinsp;\u0026plusmn;\u0026thinsp;1.25a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e42.7\u0026thinsp;\u0026plusmn;\u0026thinsp;8.1a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e91\u0026thinsp;\u0026plusmn;\u0026thinsp;16a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.47\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e11.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2abcd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e8.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e6.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eHigh spheroid\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e235.2\u0026thinsp;\u0026plusmn;\u0026thinsp;17.1c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e13.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.7ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e9.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e45.3\u0026thinsp;\u0026plusmn;\u0026thinsp;3.9b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e18.78\u0026thinsp;\u0026plusmn;\u0026thinsp;1.25a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e98.1\u0026thinsp;\u0026plusmn;\u0026thinsp;8.1c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e208\u0026thinsp;\u0026plusmn;\u0026thinsp;16c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.48\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e11.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e8.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e6.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1ab\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eOblong\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e200.5\u0026thinsp;\u0026plusmn;\u0026thinsp;13.2bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e17.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e7.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e37.2\u0026thinsp;\u0026plusmn;\u0026thinsp;3.0ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e18.36\u0026thinsp;\u0026plusmn;\u0026thinsp;0.97a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e83.4\u0026thinsp;\u0026plusmn;\u0026thinsp;6.3bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e163\u0026thinsp;\u0026plusmn;\u0026thinsp;12bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.52\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e11.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1bcd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e9.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e7.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1ab\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eOblong compressed\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e162.5\u0026thinsp;\u0026plusmn;\u0026thinsp;20.9abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e17.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e7.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e24.1\u0026thinsp;\u0026plusmn;\u0026thinsp;4.8ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e14.70\u0026thinsp;\u0026plusmn;\u0026thinsp;1.53a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e67.8\u0026thinsp;\u0026plusmn;\u0026thinsp;10.0abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e135\u0026thinsp;\u0026plusmn;\u0026thinsp;19abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.51\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e11.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2abcd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e9.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e6.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1ab\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eOblong pointed\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e160.8\u0026thinsp;\u0026plusmn;\u0026thinsp;20.9abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e19.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e31.4\u0026thinsp;\u0026plusmn;\u0026thinsp;4.8ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e19.39\u0026thinsp;\u0026plusmn;\u0026thinsp;1.53a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e58.8\u0026thinsp;\u0026plusmn;\u0026thinsp;10.0abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e124\u0026thinsp;\u0026plusmn;\u0026thinsp;19abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.48\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e11.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2abcd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e8.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e6.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eObovate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e157.3\u0026thinsp;\u0026plusmn;\u0026thinsp;13.2ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e12.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e8.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2cd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e25.4\u0026thinsp;\u0026plusmn;\u0026thinsp;3.0a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e16.56\u0026thinsp;\u0026plusmn;\u0026thinsp;0.97a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e64.0\u0026thinsp;\u0026plusmn;\u0026thinsp;6.3abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e121\u0026thinsp;\u0026plusmn;\u0026thinsp;12ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.53\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e11.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1abcd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e8.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e7.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1ab\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eOvate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e131.4\u0026thinsp;\u0026plusmn;\u0026thinsp;14.8a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e13.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e7.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e24.8\u0026thinsp;\u0026plusmn;\u0026thinsp;3.4a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e18.10\u0026thinsp;\u0026plusmn;\u0026thinsp;1.08a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e46.8\u0026thinsp;\u0026plusmn;\u0026thinsp;7.0a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e107\u0026thinsp;\u0026plusmn;\u0026thinsp;14ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.43\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e11.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e8.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e6.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1ab\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRhomboid\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e187.9\u0026thinsp;\u0026plusmn;\u0026thinsp;20.9abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e15.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9bcd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e9.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4cd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e29.0\u0026thinsp;\u0026plusmn;\u0026thinsp;4.8ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e15.54\u0026thinsp;\u0026plusmn;\u0026thinsp;1.53a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e73.2\u0026thinsp;\u0026plusmn;\u0026thinsp;10.0abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e134\u0026thinsp;\u0026plusmn;\u0026thinsp;19abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.55\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e12.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2cd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e8.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e7.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1ab\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSpheroid\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e135.1\u0026thinsp;\u0026plusmn;\u0026thinsp;17.1ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e10.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.7a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e8.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3bcd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e25.5\u0026thinsp;\u0026plusmn;\u0026thinsp;3.9a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e18.80\u0026thinsp;\u0026plusmn;\u0026thinsp;1.25a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e57.5\u0026thinsp;\u0026plusmn;\u0026thinsp;8.1ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e124\u0026thinsp;\u0026plusmn;\u0026thinsp;16ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.45\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e11.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e8.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e6.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSpheroid emarginate\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e156.8\u0026thinsp;\u0026plusmn;\u0026thinsp;29.6abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e10.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.3ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e9.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6cd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e22.0\u0026thinsp;\u0026plusmn;\u0026thinsp;6.7ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e14.07\u0026thinsp;\u0026plusmn;\u0026thinsp;2.17a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e71.8\u0026thinsp;\u0026plusmn;\u0026thinsp;14.1abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e135\u0026thinsp;\u0026plusmn;\u0026thinsp;27abc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.58\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e11.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3abcd\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e9.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e7.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2ab\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cb\u003eG/mean\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cb\u003e170.2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u003cb\u003e15.5\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u003cb\u003e8.1\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u003cb\u003e30.2\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cb\u003e17.55\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cb\u003e66.6\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cb\u003e133\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e\u003cb\u003e0.50\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e\u003cb\u003e11.7\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e\u003cb\u003e8.8\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e\u003cb\u003e7.0\u003c/b\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eFour seed colours (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) were found among the fruit shapes, including light brown (16% of all seeds), brown (46%), dark brown (35%), and greyish brown (3%).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThree seed shapes were found (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The predominant shape was reniform (55%) followed by very reniform (44%). The least shape was oblong seed (1%). All the seeds had spots in varying density on their seed coats. Light brown seed (A), dark brown seed (C) and greyish brown seed (D) were light spotted while brown seed (C) were densely spotted.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003e3.5 Principal Component Analysis of Fruit and Seed Traits\u003c/h2\u003e \u003cp\u003ePrincipal Component Analysis of the 11 standardised quantitative traits yielded three principal components (PCs) with eigenvalues\u0026thinsp;\u0026gt;\u0026thinsp;1, together explaining 77.3% of total variance. PC1 explained 42.0% of variance and was strongly loaded by overall fruit size traits: fruit weight (r\u0026thinsp;=\u0026thinsp;0.967), seed weight (r\u0026thinsp;=\u0026thinsp;0.941), pulp weight (r\u0026thinsp;=\u0026thinsp;0.861), number of seeds (r\u0026thinsp;=\u0026thinsp;0.825), and fruit width (r\u0026thinsp;=\u0026thinsp;0.793), and thus represents a latent fruit productivity axis. PC2 explained 24.4% of variance and was dominated by individual seed dimension traits seed length (r\u0026thinsp;=\u0026thinsp;0.802), seed thickness (r\u0026thinsp;=\u0026thinsp;0.742), seed width (r\u0026thinsp;=\u0026thinsp;0.739), and single seed weight (r\u0026thinsp;=\u0026thinsp;0.724) representing a seed size axis. PC3 explained 10.8% of variance and was primarily associated with pulp percentage (r\u0026thinsp;=\u0026thinsp;0.816) and fruit length (r\u0026thinsp;=\u0026thinsp;0.380), representing a pulp quality axis. The biplot (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e) showed partial separation of populations along PC1, with Karonga and Salima positioned towards the high fruit productivity end, while Chikwawa clustered at the low end. Populations overlapped considerably along PC2, indicating broadly similar individual seed dimensions across populations. Neno and Chikwawa showed wider 90% confidence ellipses, indicating greater within-population variability in overall fruit traits.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003e3.6 Tree Morphological Characteristics and Diameter at Breast Height\u003c/h2\u003e \u003cp\u003eBark colour was predominantly grey across all five districts (93.7% of trees overall), with reddish-brown bark recorded mainly in Karonga (26.7%). The most common tree shape was roundish (55.6% of all trees), followed by semi-circular (33.3%) and ellipsoid (11.1%). Mangochi was distinguished by a dominance of semi-circular canopy form (93.3%), while all sampled trees in Chikwawa were roundish (100%). Cone-shaped trunks were most prevalent across all districts (55.6%), followed by cylindrical (21.4%), concave (15.9%), and bell-shaped (7.1%). Spreading growth habit was most common overall (65.1%), though Chikwawa had a notably higher proportion of drooping trees (61.9%), and erect growth was most prevalent in Salima (26.7%) and Neno (23.3%). Trees were predominantly collected from fallow land, except in Chikwawa (95.2% on farmland) and Mangochi (50% in forest/woodland), suggesting differences in land-use and conservation context across populations (Fig.\u0026nbsp;8). DBH varied significantly among districts (F\u0026thinsp;=\u0026thinsp;12.25, p\u0026thinsp;\u0026lt;\u0026thinsp;0.001; Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Karonga had the largest trees (mean 309.2\u0026thinsp;\u0026plusmn;\u0026thinsp;16.6 cm), significantly larger than Neno (187.0\u0026thinsp;\u0026plusmn;\u0026thinsp;7.9 cm) and Mangochi (227.0\u0026thinsp;\u0026plusmn;\u0026thinsp;9.1 cm; both p\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Chikwawa (272.8\u0026thinsp;\u0026plusmn;\u0026thinsp;10.3 cm) and Salima (231.2\u0026thinsp;\u0026plusmn;\u0026thinsp;9.4 cm) were intermediate. The overall DBH range across all trees was 104\u0026ndash;422 cm (Table\u0026nbsp;7).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab5\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 5\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eDiameter at breast height (DBH) of baobab trees across five districts in Malawi. Different letters indicate significant differences (Tukey HSD, p\u0026thinsp;\u0026lt;\u0026thinsp;0.05).\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDistrict\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003en (trees)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMean\u0026thinsp;\u0026plusmn;\u0026thinsp;SE (cm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eRange (cm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eTukey\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eChikwawa\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e273.1\u0026thinsp;\u0026plusmn;\u0026thinsp;10.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e196.1\u0026ndash;347\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eab\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eKaronga\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e309\u0026thinsp;\u0026plusmn;\u0026thinsp;19.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e179.9\u0026ndash;422\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ea\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMangochi\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e226.7\u0026thinsp;\u0026plusmn;\u0026thinsp;10.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e133.1\u0026ndash;356.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ebc\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNeno\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e186.9\u0026thinsp;\u0026plusmn;\u0026thinsp;9.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e120\u0026ndash;294.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003ec\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSalima\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e231.2\u0026thinsp;\u0026plusmn;\u0026thinsp;12.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e104.1\u0026ndash;374.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eb\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"5\"\u003eF\u0026thinsp;=\u0026thinsp;12.25, p\u0026thinsp;\u0026lt;\u0026thinsp;0.001. Different letters indicate significant differences (Tukey HSD, p\u0026thinsp;\u0026lt;\u0026thinsp;0.05).\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"4.0 Discussion","content":"\u003cp\u003eThis study demonstrates substantial phenotypic variation in fruit and seed traits among wild and parkland baobab populations in Malawi, highlighting important opportunities for domestication, germplasm selection, and conservation. These differences are not merely descriptive; they point to functionally important diversity relevant to agroforestry systems where baobab contributes food, income, and resilience. This supports earlier work identifying \u003cem\u003eAdansonia digitata\u003c/em\u003e as a nutritionally important but still under-domesticated tree species for which trait characterization remains a necessary step toward improvement and wider use (Kehlenbeck et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Assogba et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Kitony et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe occurrence of 13 fruit-shape classes across five populations indicates considerable intraspecific diversity. This exceeds some earlier descriptions from southern Africa and aligns more closely with later studies elsewhere in Africa (Wickens 1982; Gebauer and Luedeling \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Gurashi and Kordofani, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). The fact that some fruit shapes observed in Malawi were not reported in Sudan further suggests that baobab morphological diversity remains incompletely documented across its range. This is important because it indicates that southern African populations should not be viewed as morphologically marginal, but rather as potentially valuable reservoirs of undercharacterized diversity.\u003c/p\u003e \u003cp\u003eA key finding was that each sampled tree consistently produced a single fruit-shape type, with no within-tree polymorphism. Although the present study did not test heritability directly, this within-tree consistency suggests that fruit shape is a relatively stable phenotype and may be under substantial genetic or maternal developmental control. For agroforestry domestication, this is encouraging because it strengthens the rationale for selecting elite mother trees and propagating them vegetatively, consistent with earlier baobab domestication studies and successful grafting work in Malawi and Kenya (Tchoundjeu et al. 2006; Anjarwalla et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Jenya et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Recent genomic evidence showing substantial population structure and adaptive diversity in baobab further supports the view that at least part of the observed variation has a biological basis (Kitony et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFruit shape was also associated with clear differences in productivity. Variation in fruit weight, pulp weight, and total seed weight shows that fruit shape is not simply a descriptive character but one linked to traits of practical value. Similar patterns have been reported in Sudan and Mali (De Smedt et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Gurashi and Kordofani, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). In this study, high spheroid and oblong fruits appear particularly promising where the objective is to maximize pulp and total seed output. However, the results also show that productivity is multidimensional. Clavate fruits produced the largest individual seeds, whereas high spheroid fruits tended to maximize total seed mass through greater seed number. This suggests that selection for pulp-oriented production and selection for seed-oriented uses may favor different phenotypes. That distinction matters in agroforestry settings, where baobab may be managed for multiple products including pulp, oil, propagation material, and nursery stock. A single ideotype is therefore unlikely to serve all production goals.\u003c/p\u003e \u003cp\u003eThe identification of four seed-coat colour classes further suggests that current baobab descriptor systems may not yet capture the full phenotypic range present in eastern and southern Africa (Kehlenbeck et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). This could have practical relevance because seed-coat colour in other species has been linked to biochemical composition and dormancy-related traits (Dixon and Sumner \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Ochuodho and Modi \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Although such relationships were not tested here, the finding points to useful follow-up work on germination and nutritional quality. By contrast, seed shape appears comparatively conserved and may be less informative than fruit shape or seed-coat colour in distinguishing superior material.\u003c/p\u003e \u003cp\u003eThe uneven distribution of fruit shapes among populations has clear conservation implications. Some shapes were widespread, whereas others were rare and locally restricted, indicating that baobab diversity in Malawi is geographically distributed rather than concentrated in a single site. Conservation strategies focused on only one or two populations would therefore risk missing part of the country\u0026rsquo;s phenotypic diversity. This supports maintaining multiple populations in situ while also sampling broadly for ex situ conservation and domestication. For agroforestry development, that is an important point, improvement efforts should not focus only on the highest-yielding sites, but should also capture rare and potentially valuable phenotypes from other locations. Because fruit shapes were sampled in a balanced way, the diversity metrics reported here should be interpreted as indicators of trait representation rather than natural abundance.\u003c/p\u003e \u003cp\u003eThe PCA reinforces the view that baobab variation is structured along more than one functional axis. One axis reflected overall fruit productivity, whereas another captured individual seed morphology. This indicates that selecting for high fruit weight will not necessarily improve all other desirable traits. Similar multivariate structuring has been reported in West African baobab populations (Assogbadjo et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; De Smedt et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). For domestication and agroforestry management, the implication is straightforward, selection criteria should be matched to the intended end use from the outset. Fruit weight may be a useful field proxy where pulp production is the priority, but seed-based value chains will require additional screening for seed traits.\u003c/p\u003e \u003cp\u003eOverall, the study supports a targeted rather than generic approach to baobab domestication in Malawi. Wild and parkland populations contain substantial phenotypic diversity, but that diversity is structured in ways that matter for use, management, and conservation. High spheroid and oblong fruits appear promising for pulp-oriented improvement, whereas phenotypes with larger individual seeds may be more suitable for seed-based products. For agroforestry development, the main lesson is that domestication goals should be defined clearly before selection begins. Future work should test the stability of these traits under common garden or clonal trials and assess whether seed-coat colour is associated with germination or biochemical differences. Integrating such phenotypic screening with emerging genomic tools would provide a stronger basis for baobab improvement and conservation in smallholder agroforestry landscapes (Kitony et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e"},{"header":"5.0 Conclusion","content":"\u003cp\u003eWild and parkland baobab populations in Malawi harbor substantial and structured phenotypic variation in fruit and seed traits, making them valuable for domestication, germplasm development, and conservation. The diversity of fruit shapes, including previously unreported forms, highlights the importance of these populations as reservoirs of under characterized variation. Because fruit shape is linked to productivity, and seed traits vary in ways relevant to different end uses, baobab improvement should rely on targeted multi-trait selection rather than a single productivity measure. The geographically distributed nature of this diversity also supports multi-site conservation and broad germplasm sampling. Together, these findings provide a practical basis for baobab domestication in Malawi and reinforce the need to conserve wild populations while advancing improvement through common garden testing and genomic tools.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was supported by the Regional Universities Forum for Capacity Building in Agriculture (RUFORUM). The authors are indebted to RUFORUM for funding this project (Grant Number: RU 2014 GRG 107), and the views expressed here are not necessarily those of RUFORUM. Above all, we would like to pay our gratitude and our respects to our late colleague Prof. Dr. Chimuleke R. Y. Munthali. After helping this research Prof. Munthali sadly passed away before we could think of publishing this work. He was a dedicated professor in the Department of Forestry \u0026amp; Environmental Management at Mzuzu University, Malawi, with a passion for research on the baobab tree.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDeclaration of Competing Interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor Contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eHerbert Jenya: conceptualization, methodology, investigation, data curation, analysis, writing original draft, writing review and editing. Owen Kachala: analysis, writing review and editing.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData Availability Statement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe data that support the findings of this study are available from the corresponding author (Herbert Jenya; [email protected]) upon reasonable request. Data include fruit and seed morphological measurements, tree diameter at breast height records, and tree morphological trait observations collected across five baobab populations in Malawi.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAbere M, Yilma Z, Tsegie T, Eshete A, Alemu A (2023) Population structure and phenological attributes of Adansonia digitata L. (baobab) in Northwestern lowland area of Ethiopia. Heliyon 9:e22571. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.heliyon.2023.e22571\u003c/span\u003e\u003cspan address=\"10.1016/j.heliyon.2023.e22571\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAnjarwalla P, Ofori D, Owino A, Matuku D, Adika W, Njogu K, Kehlenbeck K (2017) Testing different grafting methods for vegetative propagation of baobab (Adansonia digitata L.) in Kenya to assist its domestication and promote cultivation. Forests Trees and Livelihoods 26:85\u0026ndash;95. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1080/14728028.2016.1202791\u003c/span\u003e\u003cspan address=\"10.1080/14728028.2016.1202791\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAssogbadjo AE, Chadare FJ, Manda L, Sinsin B (2021) A 20-year journey through an orphan African baobab (Adansonia digitata L.) towards improved food and nutrition security in Africa. Frontiers in Sustainable Food Systems 5:675382. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3389/fsufs.2021.675382\u003c/span\u003e\u003cspan address=\"10.3389/fsufs.2021.675382\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAssogbadjo AE, Gl\u0026egrave;l\u0026egrave; Kaka\u0026iuml; R, Edon S, Kyndt T, Sinsin B (2011) Natural variation in fruit characteristics, seed germination and seedling growth of Adansonia digitata L. in Benin. New Forests 41:113\u0026ndash;125. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s11056-010-9214-z\u003c/span\u003e\u003cspan address=\"10.1007/s11056-010-9214-z\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAssogbadjo AE, Kyndt T, Sinsin B, Gheysen G, Van Damme P (2006) Patterns of genetic and morphometric diversity in baobab (Adansonia digitata): populations across different climatic zones of Benin (West Africa). Annals of Botany 97:819\u0026ndash;830. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1093/aob/mcl043\u003c/span\u003e\u003cspan address=\"10.1093/aob/mcl043\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAssogbadjo AE, Sinsin B, Codjia JTC, Van Damme P (2005) Ecological diversity and pulp, seed and kernel production of the baobab (Adansonia digitata) in Benin. Belgian Journal of Botany 138:47\u0026ndash;56\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAssogba D, Idohou R, Chirwa P, Assogbadjo AE (2022) On opportunities and challenges to conserve the African baobab under present and future climates in Benin (West Africa). Journal of Arid Environments 198:104692. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.jaridenv.2021.104692\u003c/span\u003e\u003cspan address=\"10.1016/j.jaridenv.2021.104692\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBuchmann C, Prehsler S, Hartl A, Vogl CR (2010) The importance of baobab (Adansonia digitata L.) in rural West African subsistence: suggestion of a cautionary approach to international market export of baobab fruits. Ecology of Food and Nutrition 49:145\u0026ndash;172. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1080/03670241003766014\u003c/span\u003e\u003cspan address=\"10.1080/03670241003766014\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eChl\u0026aacute;dov\u0026aacute; A, Kalousov\u0026aacute; M, Mand\u0026aacute;k B, Kehlenbeck K, Prinz K, Šm\u0026iacute;d J, Van Damme P, Lojka B (2019) Genetic diversity and structure of baobab (Adansonia digitata L.) in southeastern Kenya. Royal Society Open Science 6:190854. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1098/rsos.190854\u003c/span\u003e\u003cspan address=\"10.1098/rsos.190854\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCossam FG, Dzanja JK, Kamwana B, Olumeh DE, Mith\u0026ouml;fer D (2023) Determinants of postharvest losses along the baobab value chain in Malawi. Frontiers in Sustainable Food Systems 7:1119107. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3389/fsufs.2023.1119107\u003c/span\u003e\u003cspan address=\"10.3389/fsufs.2023.1119107\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCuni Sanchez A, De Smedt S, Haq N, Samson R (2011) Comparative study on baobab fruit morphological variation between western and south-eastern Africa: opportunities for domestication. Genetic Resources and Crop Evolution 58:1143\u0026ndash;1156. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10722-010-9647-4\u003c/span\u003e\u003cspan address=\"10.1007/s10722-010-9647-4\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCuni Sanchez A, Haq N, Assogbadjo AE (2010) Variation in baobab leaf morphology and its relation to drought tolerance. Genetic Resources and Crop Evolution 57:17\u0026ndash;25. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10722-009-9447-x\u003c/span\u003e\u003cspan address=\"10.1007/s10722-009-9447-x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ede Mendiburu F (2023) agricolae: statistical procedures for agricultural research. R package version 1.3-7. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.32614/CRAN.package.agricolae\u003c/span\u003e\u003cspan address=\"10.32614/CRAN.package.agricolae\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDe Smedt S, Alaerts K, Kouyate AM, Van Damme P, Potters G, Samson R (2011) Phenotypic variation of baobab (Adansonia digitata L.) fruit traits in Mali. Agroforestry Systems 82:87\u0026ndash;97. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10457-010-9357-0\u003c/span\u003e\u003cspan address=\"10.1007/s10457-010-9357-0\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDixon RA, Sumner LW (2003) Legume natural products: understanding and manipulating complex pathways for human and animal health. Plant Physiology 131:878\u0026ndash;885. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1104/pp.102.017319\u003c/span\u003e\u003cspan address=\"10.1104/pp.102.017319\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGebauer J, Luedeling E (2013) A note on baobab (Adansonia digitata L.) in Kordofan, Sudan. Genetic Resources and Crop Evolution 60:1587\u0026ndash;1596. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10722-013-9964-5\u003c/span\u003e\u003cspan address=\"10.1007/s10722-013-9964-5\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGruenwald J (2009) Novel botanical ingredients for beverages. Clinics in Dermatology 27:210\u0026ndash;216\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGovernment of the Republic of Malawi (2005) Food security and nutrition project: project appraisal report. Food and Agriculture Organization of the United Nations\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGruenwald J, Galizia M (2005) Adansonia digitata: market brief in the European Union for selected natural ingredients derived from native species. United Nations Conference on Trade and Development\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGurashi NA, Kordofani MA (2014) Morphological variation in fruit shapes of Adansonia digitata L. from Blue Nile and North Kordofan States. Journal of Forest Products and Industries 3:106\u0026ndash;111\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eJenya H, Munthali CRY, Mhango J (2018) Amenability of African baobab (Adansonia digitata L.) to vegetative propagation techniques. Journal of Sustainable Forestry. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1080/10549811.2018.1449123\u003c/span\u003e\u003cspan address=\"10.1080/10549811.2018.1449123\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKalinganire A, Savadogo P, Ky-Dembele C, Bush D, R\u0026aelig;bild A, Tougiani A, Muchugi A, Traore FT, Kone B (2023) Evidence of early genetic variation in survival and growth traits of baobab (Adansonia digitata L.): the way forward for domestication and breeding. Agroforestry Systems 97:1221\u0026ndash;1231. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10457-023-00837-z\u003c/span\u003e\u003cspan address=\"10.1007/s10457-023-00837-z\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKamatou GPP, Vermaak I, Viljoen AM (2011) An updated review of Adansonia digitata: a commercially important African tree. South African Journal of Botany 77:908\u0026ndash;919. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.sajb.2011.08.010\u003c/span\u003e\u003cspan address=\"10.1016/j.sajb.2011.08.010\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKehlenbeck K, Padulosi S, Alercia A (2015) Descriptors of baobab (Adansonia digitata L.). Bioversity International and World Agroforestry Centre\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKitony JK, Colt K, Abramson BW, Hartwick NT, Petrus S, Konozy EHE, Karimi N, Yant L, Michael TP (2024) Chromosome-level baobab genome illuminates its evolutionary trajectory and environmental adaptation. Nature Communications 15:8833. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1038/s41467-024-53157-w\u003c/span\u003e\u003cspan address=\"10.1038/s41467-024-53157-w\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eL\u0026ecirc; S, Josse J, Husson F (2008) FactoMineR: an R package for multivariate analysis. Journal of Statistical Software 25:1\u0026ndash;18. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.18637/jss.v025.i01\u003c/span\u003e\u003cspan address=\"10.18637/jss.v025.i01\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLuckert KM, Nemarundwe N, Gibbs L, Grundy I, Hauer D, Maruzane D, Shackleton S, Sithole J (2014) Contribution of baobab production activities to household livelihoods: a case study in Zimbabwe. In: Cunningham BA, Campbell MB, Luckert KM (eds) Bark use, management, and commerce in Africa. Advances in Economic Botany, vol 17. New York Botanical Garden Press, pp 14\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMeinhold K, Darr D (2022) Keeping up with rising (quality) demands? The transition of a wild food resource to mass market, using the example of baobab in Malawi. Frontiers in Sustainable Food Systems 6:840760. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.3389/fsufs.2022.840760\u003c/span\u003e\u003cspan address=\"10.3389/fsufs.2022.840760\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMunthali CRY (2012) Use, physiology and genetic characterisation of selected natural populations of Adansonia digitata in Malawi. Doctoral dissertation, Stellenbosch University\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMunthali CRY, Chirwa PW, Akinnifesi FK (2012) Phenotypic variation in fruit and seed morphology of Adansonia digitata L. (baobab) in five selected wild populations in Malawi. Agroforestry Systems 85:279\u0026ndash;290. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10457-012-9500-1\u003c/span\u003e\u003cspan address=\"10.1007/s10457-012-9500-1\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eNgole MA, Lusambo LP, Temu BJ (2024) Baobab trees population and economic contribution: stems density, asset value and fruits value chains in Kilolo and Iringa Urban districts, Tanzania. Trees, Forests and People 16:100519. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.tfp.2024.100519\u003c/span\u003e\u003cspan address=\"10.1016/j.tfp.2024.100519\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eOchuodho JO, Modi AT (2008) Dormancy of wild mustard (Sisymbrium capense) seeds is related to seed coat colour. Seed Science and Technology 36:46\u0026ndash;55. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.15258/sst.2008.36.1.05\u003c/span\u003e\u003cspan address=\"10.15258/sst.2008.36.1.05\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eOksanen J, Simpson G, Blanchet F, Kindt R, Legendre P, Minchin P, O\u0026rsquo;Hara R, Solymos P, Stevens M, Szoecs E, Wagner H et al (2026) vegan: community ecology package. R package version 2.7-3. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.32614/CRAN.package.vegan\u003c/span\u003e\u003cspan address=\"10.32614/CRAN.package.vegan\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eOmondi M, Rimberia FK, Wainaina CM, Mukundi JBN, Orina J, Gebauer J, Kehlenbeck K (2019) Fruit morphological diversity and productivity of baobab (Adansonia digitata L.) in coastal and lower eastern Kenya. Forests Trees and Livelihoods 28:266\u0026ndash;280. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1080/14728028.2019.1659861\u003c/span\u003e\u003cspan address=\"10.1080/14728028.2019.1659861\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePatrut A, Woodborne S, Patrut RT, Rakosy L, Lowy DA, Hall G, Von Reden KF (2018) The demise of the largest and oldest African baobabs. Nature Plants 4:423\u0026ndash;426\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eR Core Team (2025) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRashidi M, Seyif K (2007) Classification of fruit shape in cantaloupe using analysis of geometrical attributes. World Journal of Agricultural Sciences 3:735\u0026ndash;740\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSantos RC, Pires JL, Correa RX (2012) Morphological characterization of leaf, flower, fruit, and seed traits among Brazilian Theobroma L. species. Genetic Resources and Crop Evolution 59:327\u0026ndash;345\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSidib\u0026eacute; M, Williams JT (2002) Baobab: Adansonia digitata L. International Centre for Underutilised Crops, Southampton\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eStadlmayr B, Wanangwa J, Waruhiu C, Jamnadass R, Kehlenbeck K (2020) Nutritional composition of baobab (Adansonia digitata L.) fruit pulp sampled at different geographical locations in Kenya. Journal of Food Composition and Analysis 94:103617. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.jfca.2020.103617\u003c/span\u003e\u003cspan address=\"10.1016/j.jfca.2020.103617\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTchoundjeu Z, Atangana A, Asaah E, Tsobeng A, Facheux C, Foundjem D, Mbosso C, Degrande A, Sado T, Kanmegne J, Mbile P, Tabuna H, Anegbeh P, Useni M (2008) Domestication, utilization and marketing of indigenous fruit trees in West and Central Africa. In: Akinnifesi FK, Leakey RRB, Ajayi OC, Sileshi G, Tchoundjeu Z, Matakala P, Kwesiga FR (eds) Indigenous fruit trees in the tropics: domestication, utilization and commercialization. CAB International, pp 137\u0026ndash;170\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTremayne MA, Richards AJ (2000) Seed weight and seed number affect subsequent fitness in outcrossing and selfing in Primula species. New Phytologist 148:127\u0026ndash;142\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eVenter SM, Witkowski ETF (2010) Baobab (Adansonia digitata L.) density, size-class distribution and population trends between four land-use types in northern Venda, South Africa. Forest Ecology and Management 259:294\u0026ndash;300. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.foreco.2009.10.016\u003c/span\u003e\u003cspan address=\"10.1016/j.foreco.2009.10.016\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWickens GE, Lowe P (2008) The baobabs: the pachycauls of Africa, Madagascar and Australia. Springer\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"agroforestry-systems","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"agfo","sideBox":"Learn more about [Agroforestry Systems](http://link.springer.com/journal/10457)","snPcode":"10457","submissionUrl":"https://submission.nature.com/new-submission/10457/3","title":"Agroforestry Systems","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Conservation, Domestication and management, Farmland, Morphology, Selection, Superior Germplasm, Ideotypes","lastPublishedDoi":"10.21203/rs.3.rs-9317216/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9317216/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eBaobab (\u003cem\u003eAdansonia digitata\u003c/em\u003e L.) is a multipurpose fruit tree with high nutritional and economic importance across sub-Saharan Africa. We characterised fruit, seed, and tree trait diversity in five wild and parkland baobab populations in Malawi to generate baseline information for conservation and germplasm selection. A total of 135 trees were sampled. Thirteen fruit shapes were identified, with ellipsoid the most common across all populations. Eleven quantitative fruit and seed traits differed significantly among the 13 fruit shapes (p\u0026thinsp;\u0026le;\u0026thinsp;0.001). High spheroid fruits consistently showed superior productivity, with the greatest fruit weight (235\u0026thinsp;\u0026plusmn;\u0026thinsp;17 g), fruit width (9.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3 cm), pulp weight (45.3\u0026thinsp;\u0026plusmn;\u0026thinsp;3.9 g), total seed weight (98.1\u0026thinsp;\u0026plusmn;\u0026thinsp;8.1 g), and number of seeds per fruit (208\u0026thinsp;\u0026plusmn;\u0026thinsp;16). Oblong fruits ranked second for most weight-related traits. PCA identified two principal axes explaining 66.4% of the total variance. PC1 (42.0%) represented a fruit productivity gradient associated with fruit weight, seed weight, and seed number, whereas PC2 (24.4%) captured independent variation in seed size traits, including length, width, and thickness. Diameter at breast height (DBH) also varied significantly among populations (F\u0026thinsp;=\u0026thinsp;12.25, p\u0026thinsp;\u0026lt;\u0026thinsp;0.001), with Karonga supporting the largest trees and Neno the smallest. Overall, the substantial trait diversity documented, particularly in Karonga and Salima, provides a strong foundation for conservation, domestication, and selection of superior baobab germplasm in Malawi.\u003c/p\u003e","manuscriptTitle":"Trait diversity in wild and parkland baobab (Adansonia digitata L.) populations in Malawi","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-04-17 16:59:49","doi":"10.21203/rs.3.rs-9317216/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2026-04-29T10:46:20+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-04-24T23:00:59+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-04-23T18:50:35+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"160974066438547016647989753987370417665","date":"2026-04-14T09:51:15+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"230442078495367064632920061434413598952","date":"2026-04-13T08:26:26+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-04-12T09:23:22+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"115689904349791608295162143818060182587","date":"2026-04-11T08:49:20+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-04-10T04:19:40+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-04-07T16:52:22+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2026-04-04T07:02:38+00:00","index":"","fulltext":""},{"type":"submitted","content":"Agroforestry Systems","date":"2026-04-04T03:24:26+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"agroforestry-systems","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"agfo","sideBox":"Learn more about [Agroforestry Systems](http://link.springer.com/journal/10457)","snPcode":"10457","submissionUrl":"https://submission.nature.com/new-submission/10457/3","title":"Agroforestry Systems","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"6e7c470d-7048-47b3-872b-16afa29863e4","owner":[],"postedDate":"April 17th, 2026","published":true,"recentEditorialEvents":[{"type":"decision","content":"Revision requested","date":"2026-04-29T10:46:20+00:00","index":"","fulltext":""}],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-05-18T11:08:34+00:00","versionOfRecord":[],"versionCreatedAt":"2026-04-17 16:59:49","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9317216","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9317216","identity":"rs-9317216","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}