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Asimina triloba

Asimina triloba, the American papaw, pawpaw, paw paw, or paw-paw, among many regional names, is a species of small deciduous tree. It has large leaves and produces a large, yellowish-green to brown fruit.

Description
A. triloba is a large shrub or small tree growing to a height of , rarely as tall as , with trunks or more in diameter. The large leaves of pawpaw trees are clustered symmetrically at the ends of the branches, giving a distinctive imbricated appearance to the tree's foliage. The leaves of the species are simple, alternate and spirally arranged, entire, deciduous, obovate-lanceolate, long, broad, and wedge-shaped at the base, with an acute apex and an entire margin, with the midrib and primary veins prominent. The petioles are short and stout, with a prominent adaxial groove. Stipules are lacking. The expanding leaves are conduplicate, green, covered with rusty tomentum beneath, and hairy above; when fully grown they are smooth, dark green above, and paler beneath. When bruised, the leaves have a disagreeable odor similar to a green bell pepper. In autumn, the leaves are a rusty yellow, allowing pawpaw groves to be spotted from a long distance. Pawpaw flowers are perfect and protogynous, about across, rich red-purple or maroon when mature, with three sepals and six petals. They are borne singly on stout, hairy, axillary peduncles. The flowers are produced in early spring at the same time as or slightly before the new leaves appear, and have a faint fetid or yeasty smell to attract pollinators. The fruit of the pawpaw is a large, yellowish-green to brown berry, long and broad, weighing from , containing several brown or black seeds in diameter embedded in the soft, edible fruit pulp. The conspicuous fruits begin developing after the plants flower; they are initially green, maturing by September or October to green, yellowish green, or brown. When mature, the heavy fruits bend the weak branches down. Full ripening often happens only after the fruit falls naturally, thus signifying a seed dispersal strategy aimed at ground-based, rather than arboreal, mammals. • Pistils: Several, on the summit of the receptacle, projecting from the mass of stamens; ovary one-celled; stigma sessile; ovules many == Taxonomy ==
Taxonomy
Asimina triloba is a member of the Annonaceae family, commonly referred to as the "custard-apple" or "soursop" family. Annonaceae are a diverse group comprising the single largest family of the order Magnoliales. They are a tropical family consisting of 112 accepted genera with about 2,200 species spread primarily across South America, Africa, and Southeast Asia. Etymology This plant's scientific name is Asimina triloba. The genus name Asimina is adapted from the Native American (probably Miami-Illinois) name or through the French colonial . The specific epithet triloba in the species' scientific name refers to its lobed fruits (three lobes, "fructibus trifidus", in Carl Linnaeus' original description). == Distribution and habitat ==
Distribution and habitat
Asimina is the only genus native to temperate regions within the otherwise tropical and subtropical flowering plant family Annonaceae. A. triloba has the northernmost range of any species in the family, extending north into southern Ontario, Canada. Within the United States, pawpaw is native to the eastern, southern, and midwestern states, ranging from New York westward to southeastern Nebraska, southward to eastern Texas and the panhandle of Florida. Pawpaw has even been successfully cultivated as far north as Nova Scotia, Canada and in several regions of Europe and Asia. Pawpaws typically grow in USDA hardiness zones 5 to 8, though some growers have had success cultivating them in USDA hardiness zone 4b with careful site selection and protection to compensate for the shorter growing season. Within its natural habitat, pawpaw grows in slightly acidic (pH 5.5 – 7.0), well-drained soils. Pawpaws are also found along floodplains, stream banks, and shaded, nutrient-rich bottomlands, but they prefer gently elevated slopes because it has a deep-reaching taproot. While pawpaws are shade tolerant, maximum fruit yields occur under full sun conditions with some wind protection. However, germinating seedlings are extremely sensitive to full sun and require partial shading during their first one or two years. == Ecology ==
Ecology
Hybridization with other Asimina species The common pawpaw is the largest and most well known of the 13 species of the Asimina genus in North America. Of those 13, 11 prefer very warm weather and have ranges rarely extending northward of Florida or coastal Alabama. Their ranges do not overlap with Asimina triloba. One southern U.S. species, Asimina parviflora, does overlap in range with pawpaw. This species is smaller than pawpaw in both its flower and its woody growth. A. parviflora is more shrublike, rarely growing even a third as tall as pawpaw. Genomically verified hybrids of A. triloba and A. parviflora have been classified as Asimina piedmontana. Pollination Pawpaw are self-incompatible, meaning pollen cannot fertilize flowers on the same plant. Other floral features of pawpaw indicative of beetle pollination include petals that curve over the downward-pointing flower center, along with food-rich fleshy bases of the inner whorl of petals. A "pollination chamber" is thereby created at a depth that only small beetles can access during the initial female-receptive stage of floral bloom. As with other well-studied species of Annonaceae, the delay in the shift from female to male floral stage offers beetles a secure, and possibly thermogenic, residence in which not only to feed but also to mate. Receptive stigmas at their arrival, followed by pollen-shedding stamens during pollinator departure, are regarded as an early form of mutualism evolved between plants and insects that is still dominant in the most ancient lineages of flowering plants, including the Magnoliids (of which Annonaceae is the most species-rich taxonomic family). Beetles are the dominant form of pollinator ascribed for genera and species within the Annonaceae family. However, two species of genus Asimina (Asimina triloba and Asimina parviflora) bear a floral character that has given rise to an alternative hypothesis that carrion or dung flies are their effective pollinators. That floral characteristic is the dark maroon color of the petals. Hence, while no scholarly papers have documented carrion or dung flies as effective pollinators in field observations, the strength of this hypothesis has led some horticultural growers to place carrion in pawpaw orchards during the bloom time. Professional papers on genus Asimina and its species have warned of the difficulties in discerning whether insects observed on or collected from flowers are effective pollinators or merely casual and thus opportunistic visitors. A citizen science project in southern Michigan utilized natural history forms of observation, along with video and photo documentation, during a "pawpaw pollinator watch" in May 2021. Larvae and adult beetle stages of Glischrochilus quadrisignatus were also documented by the citizen project on the ground-level side of rotting fruit in a pawpaw orchard in Michigan following the fruit harvest. The fruit of pawpaw is thus recognized as having coevolved with large mammals serving as long-distance seed dispersers. The megafaunal dispersal syndrome is a common feature of some plants native to the Western Hemisphere, where a large proportion of megafauna went extinct near the end of the glacial episodes. Such fruits are now regarded as evolutionary anachronisms. Another indicator of dispersal adaptation for megafauna is that pawpaw fruit (wild types and most cultivars) tend to remain green or become blotched with brown when at peak ripeness. Mammals (other than primates) rely on olfactory rather than visual clues for discerning ripe fruit, so fruit color is no signal of ripeness for large mammals. An advantage of maintaining green fruit skin throughout the ripening process is that photosynthesis can continue during this time. Following the extinction of much of the ice age megafauna, bears would have continued dispersing pawpaw seeds in their dung. Hand carrying of fruit and seeds by humans expanding from Asia into North America would have extended the range of long-distance seed dispersal. Pennsylvania, and Michigan. Small mammals, including raccoons, gray foxes, opossums, and squirrels, assist in local movements of seeds. Interactions with animals Pawpaw defends against herbivory by producing strong-smelling natural toxins known as acetogenins. Pawpaw leaves, twigs, and bark are therefore seldom consumed by rabbits, deer, or goats, nor by insects. and mules have been seen eating the leaves in Maryland. The fruit pulp is readily consumed by mammals, including opossums, foxes, squirrels and raccoons. Larvae of the zebra swallowtail (Eurytides marcellus), a butterfly, feed exclusively on young leaves of A. triloba and various other pawpaw (Asimina) species, but do not occur in great numbers on the plants. A partially citizen-led project in Pittsburgh, Pennsylvania is working on restoring pawpaw for the purpose of encouraging return of the zebra swallowtail as well as the general restoration of riparian areas. In June 2024 a zebra swallowtail was documented laying eggs on a backyard pawpaw plant in Pittsburgh. This was the first time that this butterfly species was seen there since industrialization and river slope destruction had decimated its plant host. A local news editorial wrote, "The return of the zebra swallowtail, after 87 years, is a huge success for naturalists, conservationists and native tree lovers.... Organizations like Grow Pittsburgh, Tree Pittsburgh, the Pittsburgh Parks Conservancy and dedicated chat groups for Pittsburgh conservationists made the zebra swallowtail butterfly's restoration possible." Other insects that have evolved the ability to consume pawpaws include Talponia plummeriana, the pawpaw peduncle borer, whose larvae can be found in flowers, and Omphalocera munroei, the asimina webworm, whose larvae mostly feed upon leaves. Drosophila suzukii, the spotted wing drosophila, and Zaprionus indianus, the African fig fly, have been recorded developing within pawpaw fruit collected in Connecticut, United States, marking the first known use of a member of the Annonaceae family by D. suzukii and an expansion of the known host range of Z. indianus. Patch-forming clonal growth Pawpaw is well suited to life as an understory tree. Its large seed enables significant below-ground growth before the above-ground growth needs to access sunlight for photosynthesis. As well, the species is so shade-adapted that propagation of seedlings in nursery and landscape settings may fail if the emerging plants are not protected from direct sunlight. Pawpaws are not the first to colonize a disturbed site, but because they are capable of growing in deep shade, they can establish from seed beneath mature deciduous trees and then spread into a subcanopy patch. They may even become dominant through time by depriving native canopy trees from re-establishing via seed in a treefall gap, owing to the dense shade within a pawpaw patch. Under such circumstances, the pawpaw subcanopy becomes the forest canopy, albeit at a height half as high as the usual canopy of native trees. Accessing full sunlight, the patch is then capable of producing more fruit. Upslope expansion has been attributed to a lessening of human-set fires within forested habitats of the eastern United States. Unlike common canopy trees such as oaks and pines, pawpaw has no resistance against ground fires. So a reduction in fires has enabled pawpaw, as well as other shade-adapted native trees (including American beech and striped maple), to become more common. Pawpaw exhibits a high tolerance for intense shading, even compared with other shade-adapted species such as striped maple. Pawpaw leaves cast very heavy shade, and this chokes out seedlings and saplings of most canopy species (though not native spring herbs that benefit from the late-leafing habit of pawpaw). This helps pawpaw outcompete rival species and is a contributor to its increased abundance in forests across its range. ==Conservation==
Conservation
On a global (range-wide) scale, the common pawpaw (A. triloba) has a NatureServe global conservation rank of G5 (very common). The species is, however, listed for conservation concern in the northernmost parts of its range, owing to the happenstance of where governmental boundaries exist. In the United States, the species has an N5 (very common), but is considered a threatened species in New York, and an endangered species in New Jersey. In Canada, where the species is found only in portions of southern Ontario, it has a rank of N3 (vulnerable), and a NatureServe subnational conservation rank of S3 (vulnerable) in Ontario. The Ontario Ministry of Natural Resources has given the species a general status of "Sensitive", and its populations there are monitored. In areas in which deer populations are dense, pawpaws appear to be becoming more abundant locally, since the deer avoid them but consume seedlings of most other woody plants. Disease The genus Asimina exists on only one continent, North America. This gives the pawpaw a distinct advantage in the modern world of transcontinental commerce. In contrast to North American tree genera whose ranges extend into Europe or Asia (notably, chestnut and elm), global horticultural trade is therefore unlikely to introduce diseases that could decimate pawpaw. As for native disease, the pawpaw fares very well. There are no known disease agents (including insects) that are especially damaging. Consequently, planting of pawpaw for landscaping or fruit production requires little to no attention to disease management. Habitat restoration Pawpaw is sometimes included in ecological restoration plantings, as they have many characteristics that make them ideal for repair of riparian ecosystems. The tree's fondness of wet soil and tendency to multiply clonally to form dense and well-rooted thickets can protect against erosion and runoff. As a native species, pawpaw can be planted on river slopes for erosion control, as introduced species formerly used in the eastern United States for this purpose (such as non-native bamboo species and Amur honeysuckle) are now discouraged or prohibited because of their invasiveness. The nonexistent commercial demand of pawpaw timber also protects trees used for ecological reasons from potential future harvest. == Cultivation ==
Cultivation
Cultivation is best in hardiness zones 5-9 Commercial pawpaw production tends to be less intensive than other fruit trees. Pawpaw requires very little pesticide/herbicide, even when planted as a monoculture. Because of its long taproot, pawpaw grows well even in drier upland plantings. Cultivation of pawpaws for fruit production has attracted interest, particularly among organic growers, as a fruit with few to no pests that can successfully be grown in its native environment without pesticides. The commercial cultivation and harvesting of pawpaws is strongest in southeastern Ohio and also being explored in Kentucky In New York state, due to climate change's impact on traditional fruit crops, such as apples and peaches, farmers are looking to pawpaw as a new commercial crop.