Classification Framework of Introduced Crabapple (Malus spp.) Cultivars Based on Morphological and Numerical Traits: Insights for Germplasm Conservation and Landscape Forestry
Abstract
1. Introduction
2. Materials and Methods
2.1. Plant Materials
2.2. Trait Documentation
- Floral traits (23 indicators): inflorescence type, flower type (single, semidouble, double) (Figure 1), petal number, petal shape, petal margin, flower color, bud color, flower size, fragrance, sepal shape and margin (including eriocalyx type with hairy edge sensu Rehder 1927 [15]), pistil position, stamen fertility.
- Leaf traits (12 indicators): young leaf color, mature leaf shape (Figure 2), margin, venation, pubescence, petiole length, lamina texture.
- Fruit traits (12 indicators): shape (Figure 3), size, surface gloss, color, fruit pedicel length and orientation, calyx persistence, seed number.
- Tree and bark traits (8 indicators): plant habit (tree, shrub), crown shape, branch density, thorn presence, bark color, trunk form.
2.3. Classification Criteria
2.4. Numerical Taxonomy
3. Results
3.1. Morphological Classification Framework
3.2. Key Diagnostic Traits
- Flower color was highly reliable for further subgrouping, with little annual variation.
- Fruit traits (shape, calyx persistence, pedicel orientation) provided auxiliary criteria, especially for distinguishing cultivars with similar floral morphology.
- Leaf characters, although variable under different environments, still contributed to the recognition of specific cultivars (e.g., M. ‘Adirondack’ with narrow elliptic leaves).
3.3. Numerical Taxonomy Analysis
3.4. Concordance with International Systems
3.5. International Comparison of Classification Approaches
4. Discussion
4.1. Horticultural and Forestry Validation of Classification
4.2. Implications for Germplasm Conservation and Nursery Practice
4.3. Contribution to Breeding Programs and International Germplasm Exchange
4.4. Toward a Globally Harmonized Classification Framework
4.5. Limitations and Future Directions
5. Conclusions
Author Contributions
Funding
Data Availability Statement
Conflicts of Interest
References
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| Category | Trait No. | Trait Name | Descriptor/Measurement Method |
|---|---|---|---|
| Floral traits (23) | F1 | Inflorescence type | Corymb/umbel/solitary |
| F2 | Flower type | Single/semidouble/double | |
| F3 | Petal number | Count of petals per flower | |
| F4 | Petal shape | Ovate/obovate/elliptic | |
| F5 | Petal apex | Rounded/emarginate/acute | |
| F6 | Petal margin | Entire/wavy/irregular | |
| F7 | Petal base | Narrow/broad/clawed | |
| F8 | Flower size | Diameter (cm) | |
| F9 | Flower color (anthesis) | White/pink/red/purple | |
| F10 | Flower bud color | Greenish/pink/red/purple | |
| F11 | Color change during flowering | Stable/fading/intensifying | |
| F12 | Petal texture | Thin/leathery/papery | |
| F13 | Flower fragrance | Absent/weak/moderate/strong | |
| F14 | Sepal shape | Lanceolate/ovate/triangular | |
| F15 | Sepal reflexion at anthesis | Reflexed/upright | |
| F16 | Sepal margin | Entire/serrulate/hairy (eriocalyx) | |
| F17 | Stamen fertility | Fertile/partially sterile | |
| F18 | Anther color | Yellow/red/purple | |
| F19 | Stamen number | Few (<10)/Medium (10–20)/Many (>20) | |
| F20 | Pistil position | Superior/inferior/semi-inferior | |
| F21 | Pistil number | Single/multiple | |
| F22 | Pistil length relative to stamens | Shorter/equal/longer | |
| F23 | Blooming period | Early/mid/late season | |
| Leaf traits (12) | L1 | Young leaf color | Green/reddish/purple |
| L2 | Mature leaf shape | Ovate/elliptic/lanceolate | |
| L3 | Leaf apex | Acute/acuminate/rounded | |
| L4 | Leaf base | Cuneate/rounded/cordate | |
| L5 | Leaf margin | Entire/serrated/doubly serrated | |
| L6 | Leaf size | Length × width (cm) | |
| L7 | Leaf thickness | Thin/medium/thick | |
| L8 | Leaf texture | Leathery/papery | |
| L9 | Leaf venation | Pinnate/curved/arcuate | |
| L10 | Leaf pubescence | Absent/sparse/dense | |
| L11 | Petiole length | Length in cm | |
| L12 | Petiole pubescence | Absent/sparse/dense | |
| Fruit traits (12) | FR1 | Fruit shape | Round/oblate/conical/elliptic |
| FR2 | Fruit size | Diameter (cm) | |
| FR3 | Fruit weight | g/fruit | |
| FR4 | Fruit color (mature) | Yellow/green/red/purple | |
| FR5 | Fruit surface gloss | Glossy/dull | |
| FR6 | Fruit skin thickness | Thin/thick | |
| FR7 | Fruit pedicel length | cm | |
| FR8 | Fruit pedicel orientation | Upright/drooping/spreading | |
| FR9 | Calyx persistence | Persistent/deciduous | |
| FR10 | Seed number per fruit | Count | |
| FR11 | Fruit maturation period | Early/mid/late | |
| FR12 | Fruit persistence on tree | Short/medium/long | |
| Tree and bark traits (8) | T1 | Plant habit | Tree/shrub |
| T2 | Tree height | m | |
| T3 | Crown shape | Upright/spreading/weeping/columnar | |
| T4 | Branch density | Sparse/medium/dense | |
| T5 | Branch posture | Upright/horizontal/pendulous | |
| T6 | Presence of spines | Present/absent | |
| T7 | Bark color | Gray/brown/reddish | |
| T8 | Bark texture | Smooth/fissured/peeling |
| Group | Subgroup | No. of Cultivars | Representative Cultivars | Key Diagnostic Traits |
|---|---|---|---|---|
| I. Single Flower Group (71) | Single-white (31) | 31 | M. ‘White Cascade’, M. ‘Marry Potter’, M. ‘Weeping Madonna’, M.‘Lancelot’, M. ‘Spring Snow’, M.‘Dolgo’, M.‘Fairytail Gold’, M. ‘Red Sentinel’, M. ‘King Arthur’, M. ‘Donald Wyman’, M. ‘Adirondack’, M. ‘Molten Lava’, ‘Sugar Tyme’, M. ‘Everest’, M. ‘Red Great’, M. ‘Spring Sensation’, M. ‘Guard’, M.‘Havest Gold’, M.‘Gorgeous’, M. ‘Winter Red’, M.‘Lollipop’, M.‘Firebird’, M. ‘David’, M. ‘Professor Sprenger’, M. ‘Cinderella’, M.×zumi ‘Calocarpa’, M. ‘Golden Raindrop’, M. ‘Hydrangea’, M. ‘Snow Drift’, M. ‘Sweet SugarTyme’, M.‘Almey’ | 5 petals, white corolla, simple ovate petals, weak fragrance |
| Single-red (40) | 40 | M. ‘Prairifire’, M. ‘Indian Magic’, M. ‘Velvet Pillar’, M. ‘Royal Beauty’, M. ‘Radiant’, M.‘ Profusion’, M. ‘Liset’, M. ‘Royal Raindrop’, M. × purpurea ‘Lemoinei’, M. ‘Purple Prince’,z M. ‘Red Splendor’, M. ‘Abundance’, M. ‘John Downie’, M. ‘Lisa’, M. ‘Rudolph’, M. ‘Red Baron’, M. ‘Thunderchild’, M. ‘Centurion’, M. ‘Eleyi’, M. ‘Louisa Contort’, M. ‘Show Time’, M. ‘Makamik’, M. ‘Cardinal’, M. × purpurea ‘Neville Copeman’, M. ‘Coralburst’, M. ‘Robinson’, M. ‘May’s Delight’, M. ‘Strawberry Jelly’, M. ‘Spring Sensation’, M. ‘Candymint’, M. ‘Pink Princess’, M. ‘Hopa’, M. ‘Royal Gem’, M.‘Butterball’, M.‘Regal’, M.‘Pink Spire’, M.‘Golden Hornet’, M.‘Flame’, M. ‘Red Jade’, M. Sylvestris, M.‘Indian Summer’ | 5 petals, pink/red/purple corolla, medium fragrance, variable sepal reflexion | |
| II. Semidouble Flower Group (2) | — | 2 | M. ‘Royalty’, M. ‘Coralcole’ | 6–10 petals, pink corolla, partial stamen sterility |
| III. Double Flower Group (7) | — | 7 | M. ‘Ballet’, M. ‘Brandywine’, M. ‘Kelsey’, M. ‘Klehm’s Improved Bechtel’, M. ‘Praire Rose’, M. ‘Hillier’, M. ×micromalus ‘Furong’ | >10 petals, double corolla, strong ornamental effect, reduced fertility |
| Group/Subgroup | Petal Number | Flower Color | Fragrance | Stamen Fertility | Sepal Reflexion | Fruit and Leaf Traits (Auxiliary) |
|---|---|---|---|---|---|---|
| I. Single Flower Group | ||||||
| –Single-white (31) | 5 | White (pure to creamy) | Weak or absent | Fertile | Mostly reflexed | Fruits small (1–2 cm), leaves ovate with serrated margin |
| –Single-red (40) | 5 | Pink to deep red/purple | Weak to moderate | Fertile | Reflexed or upright | Fruits variable (1–3 cm), leaves often reddish when young |
| II. Semidouble Flower Group (2) | 6–10 | Light pink | Moderate | Partially sterile | Mostly upright | Fruits medium (2–3 cm), crown spreading |
| III. Double Flower Group (7) | >10 (often 12–20) | Pink, red, or purple | Moderate to strong | Sterile or nearly sterile | Mostly upright | Fruits rare or absent, leaves broad ovate |
| Region/System | Main Classification Basis | Limitations | This Study’s Improvements |
|---|---|---|---|
| China | Emphasis on phylogenetic background and morphological traits (flower type, fruit traits, sepal persistence). | Nomenclatural inconsistencies; over-reliance on variable traits (leaf size, fruit yield). | Broader trait dataset (55 traits) combined with numerical taxonomy; reduced synonymy. |
| Europe | Descriptive manuals; focus on ornamental traits such as flower color, fruit size, and landscape value. | Lacks hierarchical framework; descriptive but subjective; cultivars often inconsistently classified. | Established a reproducible hierarchical system with clear diagnostic characters (flower type + color). |
| North America | Horticultural orientation; classification often tied to disease resistance, growth habit, and landscape adaptability. | Practical but weak in taxonomy; cultivars grouped by utility rather than morphology. | Provides a taxonomy-oriented but still horticulturally practical framework. |
| Japan | Emphasis on aesthetic features (floral density, cultural symbolism) in classification. | Cultural-horticultural approach; limited systematic studies; descriptors not standardized. | Offers standardized descriptors aligned with ICNCP, enabling cross-cultural comparison. |
| This Study | Morphological classification (flower type and color as primary criteria) validated by numerical taxonomy (R-type and Q-type clustering). | — | Provides an integrative, reproducible, and internationally comparable framework; reference model for global harmonization. |
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He, M.; Zheng, Y.; Hu, Y.; Zhao, P.; Ji, X. Classification Framework of Introduced Crabapple (Malus spp.) Cultivars Based on Morphological and Numerical Traits: Insights for Germplasm Conservation and Landscape Forestry. Forests 2025, 16, 1792. https://doi.org/10.3390/f16121792
He M, Zheng Y, Hu Y, Zhao P, Ji X. Classification Framework of Introduced Crabapple (Malus spp.) Cultivars Based on Morphological and Numerical Traits: Insights for Germplasm Conservation and Landscape Forestry. Forests. 2025; 16(12):1792. https://doi.org/10.3390/f16121792
Chicago/Turabian StyleHe, Mei, Yutao Zheng, Yuan Hu, Pan Zhao, and Xiaofan Ji. 2025. "Classification Framework of Introduced Crabapple (Malus spp.) Cultivars Based on Morphological and Numerical Traits: Insights for Germplasm Conservation and Landscape Forestry" Forests 16, no. 12: 1792. https://doi.org/10.3390/f16121792
APA StyleHe, M., Zheng, Y., Hu, Y., Zhao, P., & Ji, X. (2025). Classification Framework of Introduced Crabapple (Malus spp.) Cultivars Based on Morphological and Numerical Traits: Insights for Germplasm Conservation and Landscape Forestry. Forests, 16(12), 1792. https://doi.org/10.3390/f16121792
