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Drupe

A drupe is a type of simple, fleshy, indehiscent fruit that develops from the mature ovary of a single carpel in a flower, characterized by an outer edible layer surrounding a single hardened endocarp, or stone, that encloses one seed.^[1] The pericarp, or fruit wall, of a drupe consists of three distinct layers: the thin outermost exocarp (skin), the fleshy mesocarp (middle layer), and the hard, woody endocarp (pit) that protects the seed.^[2] This structure distinguishes drupes from other fruit types, such as berries, which lack a stony endocarp, and pomes, which have a central core rather than a single pit.^[3] Common examples of drupes include peaches (Prunus persica), cherries (Prunus avium), plums (Prunus domestica), apricots (Prunus armeniaca), and olives (Olea europaea), all of which feature a single seed within the pit and are widely cultivated for human consumption.^[4] Other notable drupes encompass mangos (Mangifera indica) and dates (Phoenix dactylifera), where the mesocarp provides the primary edible portion.^[2] In some cases, the seed enclosed by the endocarp is edible, as in almonds (Prunus dulcis), the seeds extracted from the drupe of the almond tree.^[1] Drupes exhibit variations in form and structure adapted to different ecological roles in seed dispersal, primarily through animal consumption of the fleshy exterior.^[5] For instance, the coconut (Cocos nucifera) is a large drupe with a fibrous mesocarp, facilitating water dispersal,^[6] while the seeds of many drupes, such as cherries, contain amygdalin that can break down into cyanide, deterring seed predation.^[1] Aggregate drupes, like blackberries (Rubus fruticosus), consist of a cluster of small drupelets derived from multiple ovaries on a single receptacle, and multiple drupes, such as the osage orange (Maclura pomifera), form from fused fruits of an inflorescence.^[3] These adaptations highlight the diversity within drupes, which are significant in botany for their role in angiosperm reproduction and in agriculture as key crops contributing to global food security.^[7]

Definition and Characteristics

Botanical Definition

In angiosperms, fruits are classified based on their origin and structure, with simple fruits developing from a single pistil (comprising one or more fused carpels) within the ovary of one flower, while compound fruits arise from multiple pistils or ovaries, either aggregate (from one flower) or multiple (from an inflorescence).^[3] The position of the ovary relative to other floral parts further influences fruit morphology: a superior ovary is free and positioned above the attachment point of sepals, petals, and stamens on the receptacle, whereas an inferior ovary is embedded below this attachment.^[8] Drupes characteristically form from a superior ovary.^[9] A drupe is defined botanically as an indehiscent fruit derived from a single carpel of a superior ovary, consisting of an exocarp (often a thin skin) and mesocarp (typically fleshy, but sometimes fibrous or dry) that enclose a single seed protected by a hard, lignified endocarp (often termed a stone or pit).^[10]^[11] This structure ensures the fruit does not split open at maturity to release seeds, distinguishing it from dehiscent fruit types.^[12] The term "drupe" originates from the Latin drupa, meaning an overripe or wrinkled olive, which itself derives from the Greek dryppa referring to an olive.^[13] Linnaeus first employed "drupa" in a botanical context in his Species Plantarum (1753), using it to describe certain stone-like fruits within his system of plant classification. This usage marked the evolution of the term in 18th-century botany, building on earlier classical references to olive morphology and integrating it into the emerging taxonomic framework for angiosperm fruits.^[14]

Anatomical Structure

A drupe is an indehiscent fruit that develops from a single carpel of a superior ovary, featuring a pericarp differentiated into three distinct layers surrounding a single seed.^[8] The pericarp, the fruit wall derived from the ovary, consists of the exocarp, mesocarp, and endocarp. The exocarp forms a thin, protective outer skin that serves as the fruit's initial barrier.^[10]^[8] The mesocarp, the middle layer, is typically fleshy and varies in thickness and texture, providing the edible portion in many drupes.^[15]^[8] The endocarp, the innermost layer, is sclerenchymatous and hardens into a woody or stony structure, often enclosing a single seed.^[16]^[17] Internally, the drupe contains a single seed protected by the endocarp, with the seed featuring a thin testa (seed coat) derived from the ovule integuments, surrounding the embryo and, in some species, endosperm. In cases with endosperm, the testa adheres closely to this nutritive tissue that supports embryo development; otherwise, nutrients are stored in the cotyledons, while the embryo consists of the rudimentary plant structure.^[18]^[6] In some drupes, the endocarp exhibits variations such as porosity for gas exchange or splitting into multiple units called pyrenes, each containing a seed.^[10] These layers provide key structural adaptations for seed viability. The exocarp and mesocarp protect against desiccation and pathogen invasion through their waxy or fleshy composition, while the high lignin content in the endocarp—often exceeding that of wood—resists mechanical damage and herbivory.^[19]^[8] The mesocarp's high water content further aids in maintaining internal humidity, supporting dispersal mechanisms.^[19] Standard cross-section illustrations of a typical drupe, such as a peach, depict these layers clearly, with the exocarp as the outer skin, mesocarp as the surrounding flesh, and endocarp as the central pit.^[8]

Classification and Types

Simple Drupes

Simple drupes are monocarpellary fruits that develop from the ovary of a single pistil in one flower, typically enclosing a single seed within a stony endocarp surrounded by a fleshy mesocarp and thin exocarp.^[7]^[3] These fruits are classified as simple because they arise from a single carpel, distinguishing them from more complex forms.^[20] The pericarp layers include the outer exocarp as a protective skin, the middle mesocarp as the edible fleshy portion, and the inner endocarp as the hardened pit.^[7] Common structural variants among simple drupes involve the adherence of the mesocarp to the endocarp and its texture. Freestone drupes feature a mesocarp that separates easily from the endocarp, facilitating processing and pitting for commercial uses like canning.^[21] In contrast, clingstone drupes have a mesocarp that adheres tightly to the endocarp, providing a firmer texture suitable for fresh consumption.^[21] Additionally, the mesocarp can be fiberless and smooth, as in many peaches, or fibrous and tough, as seen in coconuts where the fibers serve non-edible purposes like cordage.^[22] Botanical examples illustrate these variations. Olives (Olea europaea) are simple drupes with a minimal mesocarp that is thin and primarily valued for oil extraction rather than direct eating.^[3] Cherries (Prunus spp.), another classic example, possess a small, juicy mesocarp that contributes to their appeal as fresh fruit.^[8]^[18] Cultivation of simple drupes has involved selective breeding for desirable traits like freestone characteristics since the 19th century, with early efforts in the United States focusing on cross-pollination and selection in Georgia to improve fruit quality and ease of handling for both fresh markets and processing.^[23] These practices enhanced commercial viability by prioritizing varieties with non-adherent pits and consistent mesocarp texture.^[24]

Aggregate and Multiple Drupes

Aggregate drupes develop from a single flower possessing multiple carpels, where each carpel matures into a small drupe known as a drupelet, collectively forming a clustered fruit structure.^[8] In this arrangement, the drupelets adhere to a shared receptacle, the enlarged floral axis that supports the ovaries, creating a cohesive yet composite fruit. For instance, the raspberry (Rubus idaeus) consists of approximately 100 drupelets, each enclosing a single seed within its own stony endocarp, while the fleshy exocarp and mesocarp portions contribute to the overall edible mass.^[25] This formation contrasts with simple drupes by multiplying the number of individual units from one flower, enhancing seed protection and dispersal potential through a larger, more attractive fruit body.^[26] Multiple drupes, in contrast, arise from an inflorescence comprising numerous flowers, each producing its own drupe that subsequently fuses externally to form a syncarpium, a unified fruit mass derived from multiple ovaries across separate blooms.^[8] The mulberry (Morus spp.), for example, exemplifies this type, as its elongated fruit aggregates numerous tiny drupes from clustered female flowers, resulting in a collective structure that resembles a blackberry but originates from distinct floral origins.^[27] Despite the multi-flower derivation, the resulting syncarpium maintains individuality in each component drupe, with no central shared receptacle as in aggregates.^[28] In both aggregate and multiple drupes, structural unity is achieved either through attachment to a common receptacle or via external fusion of the pericarps, yet each drupelet preserves its distinct anatomical features, including an individual hardened endocarp surrounding the seed and a fleshy outer layer for protection and appeal to dispersers.^[29] This modularity distinguishes compound drupes from other compound fruit types, such as follicles—which are dry and split open along one suture—or achenes, which are dry, indehiscent, and feature a thin pericarp loosely attached to the seed without a stony layer.^[30] The terminology emphasizes the drupe's fleshy nature and endocarp specialization, avoiding confusion with these drier, non-fleshy forms in botanical classification.^[3]

Specialized Forms

One specialized form sometimes classified as a drupe is the tryma, a nut-like variant characterized by a dry, often winged or fibrous exocarp and mesocarp that forms a dehiscent husk enclosing a hard endocarp containing the seed.^[31] This structure is typical in the Juglandaceae family, such as walnuts (Juglans spp.) and hickories (Carya spp.), where the outer husk splits along sutures to release the nut-like endocarp, facilitating dispersal by gravity or animals.^[32] Rarely, drupes exhibit follicle-like dehiscence, where the pericarp splits along one suture to release seeds, deviating from the typical indehiscent nature of drupes. Such variants occur in certain Simaroubaceae species, where the fruit combines drupe-like features with partial dehiscence for wind-assisted dispersal.^[33] Other modifications include drupes with multiple seeds within a single endocarp, as seen in some tropical Anacardiaceae species like Spondias (hog plums), where the woody stone may contain 1 to 5 seeds, enhancing reproductive output in resource-rich environments.^[34] Additionally, drupes derived from inferior ovaries, such as the coffee cherry (Coffea spp.) in Rubiaceae, feature a fleshy pericarp surrounding two seeds in a leathery endocarp.^[35]^[36] Evolutionarily, drupes show transitions from fleshy forms suited for animal dispersal to dry variants adapted for wind or gravity, with fossil evidence of drupaceous fruits appearing in the early Eocene (approximately 50 million years ago) in deposits like the Wagon Bed Formation, indicating diversification amid cooling climates.^[37] This shift reflects adaptations in endocarp hardening and pericarp texture, as reconstructed from nuclear phylogenies showing multiple origins of fleshy drupes from dry-fruit ancestors in families like Rosaceae.^[38]

Examples

Common Drupes

Common drupes are found across diverse plant families, showcasing the fruit type's adaptability. In the Rosaceae family, the cherry (Prunus avium and P. cerasus) is a small, pendulous drupe, typically 1-2 cm in diameter with a thin exocarp and fleshy mesocarp surrounding a single stone.^[39] The peach (Prunus persica), also from Rosaceae, features a large drupe up to 7 cm across, distinguished by its fuzzy exocarp and thick, juicy mesocarp.^[40] Plums, encompassing various Prunus species in the same family, produce smooth-skinned drupes, often 2-5 cm in length, with a firm mesocarp and single pit.^[41] The apricot (Prunus armeniaca) similarly yields a drupe 3-5 cm in diameter, with a thin exocarp, fuzzy or smooth, and a hard endocarp enclosing the edible kernel, which is the seed.^[8] Almonds (Prunus dulcis) are seeds extracted from the drupe of the almond tree, where the mesocarp dries into a leathery hull around the hard endocarp. Beyond Rosaceae, other families illustrate further diversity in drupe morphology. The mango (Mangifera indica) of the Anacardiaceae family is a large, single-seeded drupe, reaching 10-25 cm in length, with a leathery exocarp and aromatic mesocarp enclosing a flat, woody endocarp.^[42] In the Oleaceae family, the olive (Olea europaea) forms an oil-rich mesocarp in its small drupe, about 1-3 cm long, where the fleshy portion is notably high in lipids.^[43] The avocado (Persea americana) from the Lauraceae family is a berry-like drupe, 5-20 cm long, with a thick, green exocarp, buttery mesocarp, and large single seed in a thin endocarp. Dates (Phoenix dactylifera), also in the Arecaceae family like the coconut, develop as elongated drupes 3-7 cm long, with a thin exocarp, fibrous mesocarp that becomes chewy when dried, and a hard endocarp surrounding one seed. The coconut (Cocos nucifera) from the Arecaceae family exemplifies an extreme form, with its massive drupe featuring a fibrous husk serving as the outer layer, up to 30 cm long, protecting the hard endocarp and single seed. Drupes display considerable variations in color, size, and seed count, highlighting their botanical diversity. Colors range from vibrant reds and yellows in cherries and peaches to greens and browns in olives and coconuts, influenced by pigmentation in the exocarp and mesocarp.^[44] Sizes vary dramatically, from the diminutive 1 cm cherry to the substantial 30 cm coconut, adapting to different dispersal mechanisms. While most drupes contain a single seed within the endocarp, as seen in these examples, the apricot (Prunus armeniaca) in Rosaceae features a kernel inside the pit that functions as the viable seed.^[8] These representative drupes occur predominantly in temperate and tropical regions worldwide, spanning multiple plant families and contributing to floral diversity.^[45]

Economic Importance

Drupes play a significant role in global agriculture, with major crops such as peaches, cherries, plums, olives, mangos, avocados, dates, and almonds contributing substantially to food production and trade. In 2023, global peach production reached approximately 24.94 million metric tons, led by China with over 16 million tons, underscoring the fruit's importance in both fresh and processed markets.^[46]^[47] Olives represent another cornerstone, with worldwide production estimated at around 10 million tons annually as of 2023, over 95% originating from Mediterranean countries like Spain, Italy, and Greece, where about 90% of the harvest is destined for olive oil extraction.^[48]^[49] Mangos, the most produced drupe, reached over 61 million metric tons globally in 2023, primarily from India (about 25 million tons), supporting fresh markets, exports, and processing into juices and dried products.^[50] Avocados yielded around 8.5 million metric tons in 2023, with Mexico leading at over 2.5 million tons, driving demand in salads, guacamole, and oils. Dates production totaled approximately 9.5 million metric tons in 2023, topped by Egypt (1.87 million tons), valued for dried fruit snacks and religious/cultural uses. Almond production, focusing on the kernel, was about 3 million metric tons in 2023, led by the United States (over 1 million tons), essential for nuts, butters, and milk alternatives derived from the drupe seed.^[51]^[52] Nutritionally, drupes are valued for their rich content of vitamins, fiber, and antioxidants, supporting human health through dietary consumption. For instance, cherries provide about 10-12% of the daily value of vitamin C per cup, along with potassium and polyphenols that exhibit anti-inflammatory properties.^[53] Mangos offer high vitamin A and C, avocados provide healthy monounsaturated fats and fiber, dates supply potassium and iron, while almonds deliver vitamin E and magnesium. The fleshy mesocarp offers dietary fiber that aids digestion, while antioxidants like anthocyanins in many stone fruits help combat oxidative stress. However, drupe kernels can pose risks; apricot seeds contain amygdalin, a cyanogenic glycoside that breaks down into hydrogen cyanide, potentially causing toxicity if consumed in excess, with safe limits typically under three small kernels per serving.^[54]^[55] Industrially, drupes are processed into a variety of products, including jams, juices, dried fruits, and oils, fueling a multibillion-dollar sector. Olives, in particular, yield olive oil used in cooking, cosmetics, and pharmaceuticals, with historical evidence of production and trade dating back to 6000 BCE in the eastern Mediterranean, where it underpinned ancient economies through export and cultural significance.^[56] Mangos and avocados are pulped for purees and oils, dates for sweeteners, and almonds for plant-based milks, enhancing global trade. Peaches and cherries are canned or juiced for year-round availability, enhancing food security and export revenues in producing regions. Production faces challenges from pests, diseases, and climate variability, impacting yields and necessitating sustainable practices. Insect pests like the oriental fruit moth and brown rot fungus affect stone fruits, requiring integrated pest management to minimize chemical use. Climate change exacerbates issues through erratic rainfall, heat stress, and reduced chilling hours, leading to lower outputs in key areas post-2020; for example, Mediterranean olive yields dropped due to droughts, prompting adoption of drought-resistant varieties and precision irrigation in sustainable farming initiatives. Similar challenges affect tropical drupes like mangos and avocados through hurricanes and pests.^[57]^[58]^[59]^[60]

Development and Reproduction

Fruit Development

Drupe fruit development initiates with pollination, where pollen is transferred to the stigma, followed by fertilization of the ovule within the ovary, typically occurring within days of successful pollination.^[61] This event triggers the enlargement of the ovary into the pericarp, the fruit wall comprising three layers derived from the carpel: the exocarp originating from the epidermal cells, the mesocarp from the middle hypodermal and parenchyma layers, and the endocarp from the innermost cells adjacent to the seed.^[17] The process unfolds in phases, beginning with rapid cell division and expansion post-fertilization, leading to initial fruit set.^[62] Hormonal signals orchestrate these changes, with auxins and gibberellins playing key roles in promoting cell division during early ovary enlargement and subsequent cell expansion in the mesocarp.^[63] As development progresses, the endocarp undergoes lignification, hardening into a protective stone around 4-6 weeks post-bloom, a process driven by lignin deposition that completes within 2-3 weeks.^[17] Ripening is regulated by ethylene, which induces climacteric patterns in species like peaches—marked by a burst in ethylene production triggering softening and color change—contrasting with non-climacteric behavior in cherries, where abscisic acid predominates without an ethylene surge.^[63] In the peach (Prunus persica), a representative drupe, fruit development from bloom to harvest spans approximately 4-6 months, encompassing an initial 30-day rapid growth phase, a slower pit-hardening period, and a final 4-6 weeks of mesocarp expansion before maturity.^[64]^[65] Environmental factors, particularly temperature, influence stone hardening; elevated spring temperatures can accelerate early growth but lead to incomplete lignification or defects like split pits if excessive.^[66] Parthenocarpy, the development of seedless drupes without fertilization, occurs in some cultivated varieties of peaches and cherries, often induced by gibberellin applications that mimic hormonal signals for fruit set, resulting in pitless or underdeveloped-stone fruits.^[67]

Seed Dispersal and Ecology

The primary mechanism of seed dispersal for drupes is zoochory, particularly endozoochory, in which frugivorous animals consume the fleshy mesocarp and excrete the durable endocarp enclosing the seed, thereby transporting it away from the parent plant to reduce competition and predation risk. This process is highly effective for propagation, as the endocarp's lignified structure resists mechanical damage during digestion. For smaller drupes like cherries (Prunus spp.), birds such as the American robin (Turdus migratorius), cedar waxwing (Bombycilla cedrorum), and blue jay (Cyanocitta cristata) ingest the fruit and deposit seeds via feces over distances of several kilometers, promoting colonization of new habitats. Larger drupes, such as peaches (Prunus persica) and mangos (Mangifera indica), are dispersed by mammals like bears (Ursus spp.) and primates, which often consume the pulp and either excrete or discard the intact endocarp, facilitating long-distance movement in forested ecosystems.^[19]^[68] Drupes have evolved ecological adaptations to enhance animal attraction and seed viability during dispersal. The mesocarp typically ripens to bright colors (e.g., red, black, or blue) and produces volatile organic compounds (VOCs) that emit appealing odors, serving as visual and olfactory signals to frugivores indicating nutritional rewards and ripeness. Research on nocturnal mouse lemurs (Microcebus spp.) in Madagascar forests shows that fruits with high blue-yellow chromatic contrast against foliage and elevated VOC emissions (e.g., >15,500 arbitrary units) are preferentially consumed and dispersed, increasing seed removal rates by up to 40%. The endocarp's hardened, lignified layer further protects the seed from digestive acids and abrasion, with high survival rates during gut passage (often near 100% for intact endocarps), though viability can vary with disperser physiology and seed traits; this durability ensures a portion of seeds germinate post-deposition, often with scarification benefits from gastric treatment.^[69]^[19]^[70] Drupes play key roles in habitat dynamics, particularly in forest succession and supporting biodiversity across biomes. As pioneer species, aggregate drupes like those of raspberry (Rubus idaeus) rapidly colonize disturbed sites such as post-fire clearings or logged areas, peaking in abundance 2–4 years after disturbance and stabilizing soil while providing early food sources that attract wildlife, thereby facilitating the transition to mid-successional trees like birch (Betula spp.) and aspen (Populus spp.). In tropical biomes, drupes contribute to exceptional biodiversity by interacting with diverse frugivore assemblages (e.g., over 100 bird and mammal species per forest), promoting gene flow and maintaining high plant species richness through efficient zoochory in humid, low-latitude environments. Temperate biomes, by contrast, feature fewer dispersers but rely heavily on drupes for woodland regeneration, where they enhance ecosystem resilience by enabling recolonization of fragmented habitats and supporting moderate but vital biodiversity levels.^[71]^[72]^[73] Habitat loss and declining frugivore populations threaten these dispersal processes, particularly in temperate regions. Since the 1970s, North American bird populations—key dispersers for many drupes—have plummeted by nearly 3 billion individuals (a 29% decline), driven by deforestation, agricultural intensification, and climate impacts, which fragment forests and reduce seed deposition distances by up to 95% in some areas. This has cascading effects, including clustered seed shadows under parent plants, heightened predation, and diminished plant recruitment, potentially altering temperate forest composition and biodiversity.^[74]^[75]^[76]

Distinctions from Other Fruits

Comparison with Berries

Drupes and berries represent two distinct categories of simple fleshy fruits, both developing from the ovary of a single flower with superior or inferior positioning, yet they diverge markedly in pericarp composition and seed accommodation.^[77]^[30] The primary structural difference lies in the endocarp: in drupes, it is hard, lignified, and typically thick, forming a protective stony layer around a single seed or locule, while in berries, the endocarp remains thin, fleshy, and non-sclerenchymatous, enabling multiple seeds to be embedded throughout the pericarp without such rigid enclosure.^[78]^[79]^[18] Both fruit types are indehiscent, retaining their seeds internally at maturity without splitting, which supports animal-mediated dispersal.^[30] Developmentally, drupes and berries arise from fertilized ovaries, but drupes feature specialized sclerification of the inner pericarp layers to produce the characteristic pit, contrasting with the uniform fleshy maturation in berries.^[78] Misclassifications often stem from culinary usage, where "berries" denotes small, juicy fruits like strawberries—an aggregate of achenes—and raspberries—an aggregate of drupes—while botanically, true berries encompass tomatoes, grapes, and bananas due to their single-ovary origin and fleshy pericarp.^[80]^[2]^[81] This botanical-gastronomic divide underscores how everyday language prioritizes taste and appearance over anatomical criteria.^[82] From an evolutionary perspective, drupes have adapted for endozoochory through their hardened endocarp, which shields the seed from digestive breakdown during passage through vertebrate guts, whereas berries promote broader fleshy allure and multi-seed production to enhance dispersal opportunities across diverse animal vectors.^[83]^[84]

Comparison with Nuts and Other Fruit Types

Drupes differ from true nuts primarily in their pericarp structure and texture. True nuts, such as acorns (Quercus spp.), are indehiscent, single-seeded fruits with a dry, hard pericarp throughout, derived from a multi-carpellate ovary where only one seed develops, providing rigid protection without fleshy tissue.^[9] In contrast, drupes feature a differentiated pericarp: a thin exocarp, a fleshy mesocarp, and a hard, woody endocarp (the "stone") enclosing a single seed, as seen in peaches (Prunus persica).^[8] Many culinary "nuts," like almonds (Prunus dulcis), pistachios (Pistacia vera), and cashews (Anacardium occidentale), are actually drupes or "drupaceous nuts," where the edible kernel is the seed, and the surrounding husk is a fleshy precursor that dries or is removed during processing.^[4] Beyond nuts, drupes contrast with other dry fruit types in seed number, dehiscence, and pericarp characteristics. Achenes, such as those in sunflowers (Helianthus annuus), are small, indehiscent, single-seeded fruits from a single carpel, with a thin pericarp tightly fused to the seed at one point only, lacking the multilayered, stony endocarp of drupes.^[11] Capsules, exemplified by poppies (Papaver spp.), are dry, dehiscent fruits from multiple fused carpels, splitting open to release numerous seeds, unlike the indehiscent, typically single-seeded nature of drupes.^[11] Pepos, a specialized berry type in the Cucurbitaceae family like watermelons (Citrullus lanatus), arise from an inferior ovary with a leathery rind (exocarp and mesocarp fused) surrounding multi-seeded, fleshy pulp, differing from drupes in lacking a distinct hard endocarp and often containing multiple seeds.^[8] Functionally, drupes and nuts serve distinct ecological roles in seed dispersal. The fleshy mesocarp of drupes attracts animals for endozoochory, where ingestion and excretion aid seed spread, promoting dispersal over short to medium distances in forested or shrubby habitats.^[85] Nuts, being entirely dry and durable, often rely on scatter-hoarding by rodents (e.g., squirrels caching acorns) or abiotic means like gravity and wind, enabling longer-distance dispersal in open or woodland ecosystems while resisting decay.^[86] Historical botanical reclassifications have clarified overlaps, particularly for nut-like drupes. In 20th-century taxonomy, walnuts (Juglans spp.) and hickory nuts (Carya spp.) were reclassified from true nuts to trymae—drupaceous fruits with a dehiscent outer husk enclosing a nut-like inner shell—reflecting their accessory tissue origins beyond the ovary.^[87] In the APG IV classification system (2016), drupes and nuts appear across diverse angiosperm orders, underscoring their convergent evolution as indehiscent, single-seeded forms adapted to various clades, such as Rosales for drupes in Rosaceae and Fagales for nuts in Fagaceae.

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Apr 25, 2022 · We have clingstone peaches, freestone peaches, and even semi-freestone peaches. All three can be found as white or yellow peaches.Missing: drupes | Show results with:drupes
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https://bio.libretexts.org/Bookshelves/Botany/The_Science_of_Plants_-_Understanding_Plants_and_How_They_Grow_(Michaels_et_al.)/08:_Fruit/8.01:_Fruit_Morphology
[23]
How a Navajo Scientist Is Helping to Restore Traditional Peach ...
Jun 22, 2022 · U.S. experimentation with cross-pollinating, selecting, and budding peach varieties originated in the mid-1800s in the State of Georgia. The ...
[24]
https://extension.psu.edu/fruit-breeding-peach-breeding-to-improve-fruit-quality
[25]
Raspberries : USDA ARS
Jun 26, 2018 · Fruits of raspberries and blackberries are compound drupes composed of small turgid drupelets, each of which contains 1-2 seeds. Wild species ...
[26]
3. Botany | NC State Extension Publications
Feb 1, 2022 · Cherries, nectarines, peaches, and plums are drupes with a single hard seed in the center (Figure 3–40). Apples, pears, and quince are known as ...
[27]
[PDF] Morus rubra L., red mulberry - Michigan Natural Features Inventory
The fruits, which are animal-dispersed, are multiples of tiny drupes (a drupe being a fleshy fruit with a stone-like seed) consolidated into blackberry-like ...
[28]
Morus alba (White Mulberry)
Fertilized flowers on female trees produce sweet, edible blackberry-like fruits (cylindrical drupes to 1inch long) that mature in June. Fruits ripen to white or ...
[29]
Fruit Types | Biology 343 – Plants and People - UBC Blogs
Examples: cherry, peach, plum. The small drupes that form the aggregate fruits of raspberries and blackberries are called drupelets.
[30]
8.1: Fruit Morphology - Biology LibreTexts
Jul 27, 2022 · The fibrous mesocarp is used for other purposes like floor coverings in high-foot-traffic areas where a tough, durable fiber is needed. Cross ...
[31]
Juglandaceae - FNA - Flora of North America
Nov 5, 2020 · Fruits large nuts [or samaras], nuts enclosed in dehiscent or indehiscent, fibrous-fleshy or hard involucres (husks), thus ± drupelike.
[32]
Simaroubaceae | Fruit and Seed Family ID - IDtools
an indehiscent, fleshy fruit with one or a few to many seeds. The flesh may be homogenous throughout. Or, if the outer part is hard, firm, or leathery, referred ...
[33]
HS63/MG059: Spondias Growing in the Florida Home Landscape
The fruit of all three species are oval to oblong drupes, possessing a stone (woody endocarp) that may have 1 to 5 seeds. Dry season determines when ...
[34]
CHAPTER 1: Introduction to Coffee Plant and Genetics - Books
Jan 14, 2019 · The fruit of the coffee plant is typically described as a drupe: a fleshy, indehiscent fruit with a pericarp that is clearly differentiated into ...
[35]
Rubiaceae -- the madder or coffee family (450-606/ 6500
Rubiaceae -- the madder or coffee family (450-606/ 6500 ... inferior ovary; 2 locules with many axile ovules/locule; 1 style, undivided. Fruit= berry, drupe ...
[36]
[PDF] the early middle eocene wagon bed carpoflora of central wyoming ...
Within the drupaceous forms, individual units can involve either a single fruit or several fruits fused into “polydrupes”. (Stone et al. 1998, Callmander et al.
[37]
Evolution of Rosaceae Fruit Types Based on Nuclear Phylogeny in ...
Ancestral character reconstruction for fruit types supports independent origins of fleshy fruits from dry-fruit ancestors, including the evolution of drupes ( ...
[38]
Prunus avium (Sweet Cherry)
The fruit is a fleshy drupe, yellow or red with a large pit (stone). It has occasionally escaped from gardens and naturalized in the landscape.
[39]
[PDF] What Are Stone Fruits
They come in clingstone and freestone varieties, which differ in how easily the flesh separates from the pit. Nectarines: Very similar to peaches but with ...
[40]
Prunus cerasifera - North Carolina Extension Gardener Plant Toolbox
The fruit is a smooth, round, yellow, or red drupe that is up to 1.25-inches in diameter and ripens mid to late summer. Cultivars may be reddish-purple fruits.
[41]
HS2/MG216: Mango Growing in the Florida Home Landscape
The fruits possess a single large, flattened, kidney-shaped seed that is enclosed in a woody husk. Seed Types. Mango varieties produce either monoembryonic ...
[42]
[PDF] Olive Trees Olives, Olive Oil (Olea Europaea )
The olive fruit is a drupe, spherical or elliptic in shape and consists of the exocarp (skin), which contains stomata, the mesocarp (flesh), which is the edible.
[43]
Tropical fruits & subtropical fruits
The fruit is a fleshy drupe with a thick yellowish-red skin and a large seed. The size, shape, and quality mangos vary greatly. The lengths are 3-5 in. The pulp ...
[44]
[PDF] Woody Plants at Jasper Ridge, by Toni Corelli - Stanford University
Some examples are: coffee, mango, olive, almond, apricot, cherry, nectarine, peach, pistachio, plum, most palms (including date, coconut and oil palms). See.
[45]
Peaches and Nectarines Market - Size, Share & Industry Analysis
Mar 28, 2025 · Data from the USDA indicates that global production of peaches and nectarines climbed to 24.94 million metric tons in 2023, up from 24.05 ...
[46]
Which Country is the Largest Producer of Peaches in the World?
Jun 27, 2025 · China ranks first in global peach production, yielding approximately 16,800,000 tonnes each year. ... Note: Figures are based on FAO 2023 ...<|control11|><|separator|>
[47]
World Olive Production by Country - AtlasBig.com
Spain is the largest olive producer in the world with 8,256,550 tonnes production per year. Greece comes second with 3,240,063 tonnes yearly production.
[48]
https://www.atlasbig.com/countries-olive-production
[49]
7 Impressive Health Benefits of Cherries - Healthline
Dec 4, 2024 · Vitamin C: 12% of the daily value (DV); Potassium: 10% of the DV; Copper: 10% of the DV; Manganese: 5% of the DV; Vitamin B6: 0.08 milligrams ( ...
[50]
A Review of the Health Benefits of Cherries - PMC - NIH
Mar 17, 2018 · Cherries are a rich source of polyphenols and vitamin C which have anti-oxidant and anti-inflammatory properties.<|control11|><|separator|>
[51]
Apricot kernels pose risk of cyanide poisoning - EFSA
Apr 27, 2016 · Eating more than three small raw apricot kernels, or less than half of one large kernel, in a serving can exceed safe levels.
[52]
The European market potential for table olives - CBI
Feb 6, 2025 · Less than 10% of all harvested olives are used as table olives. More than 90% of the remaining olives are processed into olive oil. The ...Missing: percentage | Show results with:percentage
[53]
How Many Olive Varieties Are There and Which Are the Most Popular?
Jul 30, 2020 · Today, olives are grown in dozens of countries, with 90 percent of the harvest destined for oil production and the remaining 10 percent ...<|control11|><|separator|>
[54]
https://pmc.ncbi.nlm.nih.gov/articles/PMC5872786/
[55]
Climate Change and Perennial Fruit and Nut Production
This publication explores climate-related challenges to perennial fruit and nut production, including drought, frost, flooding, and extreme weather events.
[56]
Sustainable ways to reduce the use of pesticides in pome and stone ...
Alternative methods reduce the use of pesticides in pome and stone fruits and support the productivity of the sector in a sustainable way.
[57]
From the Olive Flower to the Drupe: Flower Types, Pollination, Self ...
Apr 10, 2013 · Although the tree development directs the olive tree yield we focus here to describe the main phases, stages and key steps from blossoming to ...
[58]
Fruit Development - an overview | ScienceDirect Topics
Fruit development involves three basic phases. The first phase is the development of the ovary and the initiation of the cell division, together called a fruit ...
[59]
Phytohormones in fruit development and maturation - Fenn - 2021
Dec 4, 2020 · In both tomato and peach, auxin and ethylene increase in proportion to one another, and auxin has been shown to induce ethylene biosynthesis via ...
[60]
Peach: Fruit Development: Why Is It Important? - UC IPM
Jun 24, 2016 · The final stage, which usually begins 4 to 6 weeks before harvest, is a period of rapid growth of the skin (exocarp) and flesh (mesocarp) of the ...
[61]
Bloom and Harvest Dates, Fruit Quality Attributes, and Yield of ...
The periods between bloom and harvest for these cultivars on average were 160, 163, 163, 168, 171, 173, and 177 days, respectively, and these cultivars often ...<|separator|>
[62]
Influence of Temperature on the Development of Peach Fruit in a ...
Jan 7, 2019 · The low biomass accumulation of the fruit was related to the hardening of the endocarp with consequent end of growth, similar to the behavior ...
[63]
Options for the generation of seedless cherry, the ultimate snacking ...
Sep 28, 2022 · In some horticultural crops, fruit set and development can occur without the fertilization of the ovules in a process known as parthenocarpy ( ...
[64]
Prunus serotina - Forest Service
Long-distance seed dispersal by birds is important in the establishment of black cherry along fence rows and into forest openings, old fields, and pine ...
[65]
Colour and odour drive fruit selection and seed dispersal by mouse ...
Aug 13, 2013 · We show that both fruit colour and odour significantly predict fruit consumption and seed dispersal by Microcebus ravelobensis and M. murinus.Missing: drupe attraction
[66]
Effect of seed passage through vertebrate frugivores' guts on ...
It was found that seed dispersers commonly have an effect on the germinability of seeds, or on the rate of germination, or both, in about 50% of the plants ...
[67]
SPECIES: Rubus idaeus - USDA Forest Service
It is generally considered a pioneer or early seral species ... In many northern black spruce forests, red raspberry is present only in early successional stages ...Missing: drupe | Show results with:drupe
[68]
[PDF] Determinants of Geographic Patterns in Seed Dispersal Modes in ...
Apr 24, 2025 · 1) Geographic Trends: Zoochory is more common in warm, humid, low-latitude forests, while anemochory dominates in cooler, drier, and more open ...<|control11|><|separator|>
[69]
Seed dispersal phenology and dispersal syndromes in a subtropical ...
Zoochory was the principal dispersal mode of plants in subtropical forest understory. Dry seasons favor seed dispersal by animal and wind. Our study will ...
[70]
Nearly 3 Billion Birds Gone - Cornell Lab of Ornithology
The North American bird population is down by 2.9 billion breeding adults, with devastating losses among birds in every biome.
[71]
Ecosystem consequences of bird declines - PMC - NIH
Dec 28, 2004 · Important ecosystem processes, particularly decomposition, pollination, and seed dispersal, will likely decline as a result.
[72]
Lost birds and mammals spell doom for some plants - Rice News
Jan 12, 2022 · “We found regions where climate-tracking seed dispersal declined by 95%, even though they'd lost only a few percent of their mammal and bird ...Missing: drupe | Show results with:drupe
[73]
Fruits - Digital Atlas of Ancient Life
Sep 19, 2019 · Coconuts (Cocos nucifera) are drupes with a fibrous mesocarp. The mesocarp is often removed from fresh coconuts sold in grocery stores. The ...Missing: fiberless | Show results with:fiberless
[74]
Simple fleshy fruits : baccate,drupaceous - Berry, Drupe ... - BrainKart
Feb 10, 2016 · Drupe: This is normally a one-seeded fruit. In these fruits the pericarp is differentiated into an outer skinny epicarp, a middle fleshy and ...
[75]
5.3: Fruits - Biology LibreTexts
Jan 30, 2024 · Drupes usually have a single seed. Examples of drupes include mangoes, peaches, nectarines, and apricots. Mango dissection.png Figure 5 . 3 ...
[76]
Bananas are Berries. Raspberries are Not. - McGill University
Dec 6, 2017 · It turns out berry is actually a botanical term, not a common English one. Blackberries, mulberries, and raspberries are not berries at all, but bananas, ...Missing: culinary | Show results with:culinary
[77]
What makes a berry a berry? | Napa Master Gardener Column
Aug 31, 2024 · Raspberries and blackberries are made up of multiple drupes, each drupe developing from an ovary and containing a seed. A strawberry is also an ...
[78]
Insider's Viewpoint: True Berries vs Berry-Like - Have A Plant
There are “true” berries (avocados, barberries, currants, gooseberries, grapes, pineapple and tomatoes) and “false” berries such as blueberries, cranberries and ...
[79]
[PDF] Repeated evolution of net venation and fleshy fruits among ...
Jun 28, 2005 · 'Fleshy fruits' include berries, drupes and seeds with showy and massive arils dispersed by vertebrates, all with moist, sweet. (or oily) tissue ...
[80]
Seed Dispersal by Animals: A Role in Angiosperm Diversification?
evolution of fleshy diaspores (and endozoochorous dispersal) apparently has not constituted a sufficiently important evolutionary breakthrough to promote.
[81]
Fruit Characteristics, Life Forms, and Species Richness in the Plant ...
The average number of species in the 427 investigated genera was 23.2 (SD = 59.9).
[82]
[PDF] Seeds and Fruits - PLB Lab Websites
single carpel, the drupe is usually one-seeded. It has a hard endocarpconsisting of thick- walled sclereids (see Chapter 4), and a thin exocarp forms the skin. ...
[83]
[PDF] characterizing pecan (carya illinoinensis) germplasm - OAKTrust
Some botanists classify pecans as a true nut, while others use a more technical term called a “tryma”, which is another way to describe a drupe with a dehiscent ...Missing: botanical | Show results with:botanical