Overview

Comprehensive Description

Comments

Mayapple is a familiar woodland plant with interesting foliage. The flowers are large and attractive, but they are sparingly produced and mostly hidden by the large leaves. Mayapple develops very quickly during the warmer days of spring. There is no other plant within the state that resembles it; the only other species in this genus occurs in Asia. People can eat the ripe berries in limited amounts, even though they may be mildly toxic. The flavor is bland and resembles an overripe melon.
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Description

This herbaceous perennial plant is 1–1½' tall. Infertile plants are unbranched, producing a single leaf from a long stalk, while fertile plants produce a pair of leaves on long petioles at the apex of this stalk. The stalks are light green to pale reddish green, glabrous, and terete (circular in circumference). The leaves of infertile plants are up to 1' long and 1' across; they are orbicular in outline, fully peltate, and deeply divided in 6-9 palmate lobes. The leaves of fertile plants are similar, although they are less orbicular in outline, only marginally peltate, and they tend to have fewer lobes (typically 5-6). The leaves of both infertile and fertile plants have lobes that are obovate in shape. The outer margins of these lobes are coarsely dentate and often shallowly cleft; less typically, they are coarsely crenate, slightly undulate, or smooth (entire). The upper leaf surface is medium to dark green and glabrous. On fertile plants, the ascending petioles of the leaves are 3-6" long, light green to pale reddish green, glabrous, and terete. The petioles join the leaf blades toward the inner margins of the latter. Each fertile plant produces a single nodding flower where the 2 petioles branch from each other. This flower is about 1½" across, consisting of 6-9 white petals, 6 light green sepals, 12-18 stamens, and a superior ovary with a dome-shaped cluster of stigmata at its apex. Both the petals and sepals are oval-obovate in shape; the latter are glabrous and early-deciduous. The ovary is ovoid in shape and light green to pale yellow. The stamens have white filaments and yellow anthers. The pedicel of the flower is about 1½" long, light green to yellowish green, and glabrous. The blooming period occurs from mid- to late spring, lasting about 2-3 weeks. Individual flowers are short-lived; they have a pleasant fragrance. Each flower is replaced by an ovoid berry that is fleshy and contains several seeds. At maturity, this berry is about 1½" long and pale yellow. A berry is produced only when cross-pollination of the flower occurs. The root system is long-rhizomatous and fibrous. Mayapple often produces dense colonial colonies that exclude other spring-flowering plants.
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Distribution

National Distribution

Canada

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

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Global Range: Eastern North America, from Nova Scotia and Ontario (USDA-NRCS 1999), New England (Brumback and Mehrhoff 1996), west to Minnesota, eastern Kansas (Freeman pers. comm.) and Nebraska (Steinauer pers. comm.); south to Texas, Louisiana (USDA-NRCS 1999), throughout Alabama to the panhandle of Florida (Schotz pers. comm.); east to the Carolinas.

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Ont., Que.; Ala., Ark., Conn., Del., D.C., Fla., Ga., Ill., Ind., Iowa, Kans., Ky., La., Md., Mass., Mich., Minn., Miss., Mo., Nebr., N.H., N.J., N.Y., N.C., Ohio, Okla., Pa., R.I., S.C., Tenn., Tex., Vt., Va., W.Va., Wis.
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Physical Description

Morphology

Description

Rhizomes: annual elongation increments (2-)6-20 cm. Leaves of nonflowering shoots 2-5 dm; blade 18-38 × 18-38 cm. Flowering shoots 3-6 dm; leaves nearly opposite, slightly unequal in size; petioles 5-15 cm; proximal blades 10-35 × 14-40 cm, distal blades 6-25 × 10-33 cm. Leaf blades 5-7(-9)-parted, parts lobed or not (frequently 2-lobed), margins entire or coarsely dentate, teeth apiculate; surfaces abaxially sparsely pubescent to glabrous. Flowers solitary, nodding, fragrant; peduncle arising from angle between petioles, 1.5-6 cm; sepals orbiculate, 10-18 × 10-18 mm; petals white, rarely pink, obovate, 15-35 × 10-25 mm; stamens 2 times number of petals, 8-13 mm; filaments 3-5 mm; anthers 5-8 × 1-1.5 mm; ovaries 6-12 × 4-8 mm; style 1-2 mm; stigmas 3-6 mm. Berries yellow, rarely orange or maroon, 3.5-5.5 × 2.0-4 cm. Seeds 30-50, ovoid, 6-8 × 4-6 mm. 2 n = 12.
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Ecology

Habitat

Comments: This species is found in rich cove forests and mesic hardwood forests, typically those with soils that are not too acidic (Schafale pers. comm.); on circumneutral soils, this species can occur in dry-mesic oak-hickory forests (Schafale pers. comm.). This species is most typically found at low topographic positions (Pittillo pers. comm., Schafale pers. comm.), but it can also be found along ridgetops, especially on amphibolite substrate (Schafale pers. comm.). In northern portions of its range it is associated with maple woodlands and forests, occasionally along floodplains (Labrecque pers. comm.). It is also found in disturbed areas and along roadways (Enser pers. comm.), and persists in old gardens (Labrecque pers. comm.).

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Mixed deciduous forest, fields, moist road banks, river banks; 50-800m.
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Associations

Faunal Associations

The flowers are cross-pollinated by bumblebees and other long-tongued bees. These insects collect pollen and possibly suck nectar. The larvae of a sawfly, Aglaostigma quattuordecimpunctatum, feed on the leaves of Mayapple (Smith, 2006). Adults of a thrips, Ctenothrips bridwelli, have been found on the foliage (Stannard, 1968). The foliage of Mayapple is avoided by mammalian herbivores because of its poisonous qualities and bitter taste. The seeds and rhizomes are also poisonous. The berries are edible if they are fully ripe; they are eaten by box turtles and possibly by such mammals as opossums, raccoons, and skunks. The seeds are distributed to new locations in the feces of these animals. Photographic Location
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Flower-Visiting Insects of Mayapple in Illinois

Podophyllum peltatum (Mayapple)
(Long-tongued bees suck nectar, collect pollen, or explore the flowers; observations are from Robertson and Motten; Robertson assumed that the flowers produced nectar, but Motten states that the flowers are nectarless)

Bees (long-tongued)
Apidae (Apinae): Apis mellifera cp (Rb); Apidae (Bombini): Bombus bimaculatus cp/exp (Mtt), Bombus griseocallis sn (Rb), Bombus impatiens sn (Rb), Bombus pensylvanica sn (Rb); Anthophoridae (Eucerini): Synhalonia speciosa sn (Rb); Anthophoridae (Xylocopini): Xylocopa virginica cp/exp (Mtt)

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In Great Britain and/or Ireland:
Foodplant / spot causer
epiphyllous, rather scattered pycnidium of Phomopsis coelomycetous anamorph of Phomopsis podophylli causes spots on fading leaf of Podophyllum peltatum

Foodplant / spot causer
amphigenous, gregarious, usually on the veins pycnidium of Phyllosticta coelomycetous anamorph of Phyllosticta podophylli causes spots on live leaf of Podophyllum peltatum
Remarks: season: 9

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Population Biology

Number of Occurrences

Note: For many non-migratory species, occurrences are roughly equivalent to populations.

Estimated Number of Occurrences: 81 to >300

Comments: Probably tens or hundreds of thousands of populations are extant rangewide. Connecticut: 1?; Iowa: several dozen to hundreds; Indiana: thousands; Kansas: >100; Maryland: hundreds; North Carolina: thousands; Nebraska: 25-50?, 2 known; New Hampshire: 1?; Rhode Island: 2 (introduced); South Carolina: hundreds; Tennessee: several hundred or perhaps thousands; Vermont: 2; Quebec: 7 (5 likely introduced by Native Americans, 2 in old gardens) (Brumback and Mehrhoff 1996, APSU 1999).

Since this is such a common species throughout much of its range, these numbers can only be estimates. Additional information on species distribution and the number of populations can be gleaned from county occurrence dot maps (USDA-NRCS 1999).

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General Ecology

It is possible that North Carolina populations seldom set fruit, and that reproduction from seed may be rare (Schafale pers. comm.). Only rarely have fruit been observed in North Carolina, in contrast with abundant ripe fruit observed in southern Illinois (Schafale pers. comm.).

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Life History and Behavior

Cyclicity

Flowering/Fruiting

Flowering spring, fruiting late spring-summer; summer deciduous.
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Molecular Biology and Genetics

Molecular Biology

Barcode data: Podophyllum peltatum

The following is a representative barcode sequence, the centroid of all available sequences for this species.


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Statistics of barcoding coverage: Podophyllum peltatum

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 2
Specimens with Barcodes: 3
Species With Barcodes: 1
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Conservation

Conservation Status

National NatureServe Conservation Status

Canada

Rounded National Status Rank: N5 - Secure

United States

Rounded National Status Rank: N5 - Secure

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NatureServe Conservation Status

Rounded Global Status Rank: G5 - Secure

Reasons: This species has a very broad range, has a wide variety of wooded habitats across its range, and is regularly encountered in appropriate habitat (Schafale pers. comm.). It tolerates moderate disturbance (Homoya pers. comm.) and can be found in regenerating woodlands. At present, collection for the plant trade seems to only be at very low levels.

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Threats

Comments: There is evidence from a reliable source that plant collecting from wild populations is occurring for the plant trade on national forest lands in North Carolina (Kauffman pers. comm., Corbin pers. comm.). Collection permits for Pisgah and Nantahala National Forests in North Carolina were for the following amounts (dry weight): 1997 - 3600 lbs.; 1998 - 600 lbs.; 1999 - 3200 lbs. (Kauffman pers. comm.).

A large dealer in herbs based in the southern Appalachians sold 2,000-3,000 lbs. (dry) in 1999 (Fletcher pers. comm.).

This species is not among the high-price medicinal species in the black market (Corbin pers. comm.), and wildcrafting of this species may be very limited in scale (Suggs pers. comm.). The market has been erratic the past few years (Blakley pers. comm.), with some batches being thrown out when prices plummeted after it was announced that medications developed from this species may be carcinogenic (Fletcher pers. comm., Suggs pers. comm.). Nevertheless, there may be renewed interest in this species for other purposes (Fletcher pers. comm.), as evidenced by the collection permits within two Forest Service districts in North Carolina. There are varying reports of current sources of the bioactive compound, podophyllin, found in this species; Suggs (pers. comm.) thinks that it is from an oriental species of Podophyllum, whereas Blakley (pers. comm.) thinks it is being generated synthetically. Some companies have discontinued offering this product (Halvorsen pers. comm.). In Tennessee, this plant is collected from the wild and sold as nursery stock (Warren Co. Nursery).

A person knowledgable about the herbal medicinal trade says that the plant receives only very modest usage, estimated at less than one thousand pounds per year (M. McGuffin pers. comm.).

As with all native forest herbs, habitat conversion and urban/rural development are significant direct threats (Homoya pers. comm., Pittman pers. comm., Kunsman pers. comm., Pearson pers. comm., Schafale pers. comm.).

Threats to the two native populations in Vermont are extensive logging (although selective logging in the winter would be less damaging) and road widening (Vermont Nongame and Natural Heritage).

It is possible that North Carolina populations seldom set fruit, and that reproduction from seed may be rare (Schafale pers. comm.). Only rarely have fruit been observed in North Carolina, in contrast with abundant ripe fruit observed in southern Illinois (Schafale pers. comm.).

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Management

Biological Research Needs: There is a need to determine sustainable collection levels from healthy, wild populations (Kauffman pers. comm.).

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Relevance to Humans and Ecosystems

Benefits

Cultivation

The preference is dappled sunlight to light shade, moist to slightly dry conditions, and a rich loamy soil with abundant organic matter. This plant is easy to start from rhizomes and it will readily adapt to garden areas near deciduous trees. It is a strong colonizer and may spread aggressively in some situations. Young foliage is vulnerable to late-frost damage. The mature foliage dies down by the end of summer. Range & Habitat
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Economic Uses

Uses: MEDICINE/DRUG

Production Methods: Wild-harvested

Comments: The Podophyllum used pharmaceutically is the Indian mayapple (McGuffin pers. comm.).

Prices for this species were found as follows:

Central Tennessee, nursery: $0.40/bare root whole plant (wild-collected and sold in bundles of 50)

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Wikipedia

Podophyllum peltatum

Podophyllum peltatum, commonly called Mayapple, or May Apple,[1] (or hogapple, Indian apple, mayflower, umbrella plant, wild lemon (flavor of the fruit), wild mandrake, American mandrake (shape of rhizomes) or "devil's apple" (used for Solanum linnaeanum elsewhere)), is a herbaceous perennial plant in the family Berberidaceae, native to deciduous forests in of eastern North America.[2] Like many other spring ephemerals, it emerges from below ground before the canopy of the forest opens, and then slowly withers later in the summer; the foliage is, however, somewhat more long-lived than other spring ephemerals such as Trillium.

The stems grow to 30–40 cm (about 12-15 inches) tall, with 2 or occasionally 3 palmately lobed leaves up to 20–30 cm (about 8-12 inches) diameter with 5-9 deeply cut lobes on reproductive individuals, or one peltate (umbrella-like) leaf on sterile individuals. The single secund white flower 3–5 cm (1-2 inches) diameter, with six (rarely up to nine) petals, is produced at the axil of the two leaves (the upper two in a three-leaved plant); the flower matures into a yellow-greenish fruit 2–5 cm long.[3] The plant is widespread and appears in clonal colonies in open mesic woodlands. Individual shoots are often connected by systems of thick rhizomes.[1] As with many kinds of wild plants, the flower provides sexual reproduction while the rhizome provides asexual reproduction.[4] The former provides long distance dispersal, while the latter allows the formation of dense circular clones. There are costs to producing flowers, since the production of a flower and fruit reduces the probability that the plant will survive, or flower, in following years.[5]

Many species of plants have mycorrhizae to assist with nutrient uptake in infertile conditions.[4] Mayapple plants are considered obligately dependent upon such mycorrhizae, although it may also be facultatively dependent upon rhizome age and soil nutrient levels.[6] Plants are commonly found infected by the rust Allodus podophylli, appearing as honeycomb-patterned orange colonies under the leaves, and yellowish lesions on the upper surface.[7] [8]

Though the common name is mayapple,[9] it is the flower that appears in early May, not the "apple". The fruit or "apple" is produced early summer and ripens later in summer.

Toxicity[edit]

The ripened fruit is edible in little amounts, though when consumed in large amounts the fruit is poisonous. The rhizome, foliage and roots are also poisonous.[10] Mayapple contains podophyllotoxin,[11] which is highly toxic if consumed, but can be used as a topical medicine.

Medicinal use[edit]

Mayapple has been used by American Indians as an emetic, cathartic, and antihelmintic agent.[citation needed] They also boiled the poisonous root, and used the water to cure stomach aches.[citation needed] The rhizome of the mayapple has been used for a variety of medicinal purposes, originally by indigenous inhabitants and later by other settlers.[3]

Mayapple can be also used topically for warts, and two of its derivatives, etoposide and teniposide, have shown promise in treating some malignant neoplasms.[12][13]

Gallery[edit]

References[edit]

  1. ^ a b Gleason, H.A. 1968. The New Britton and Brown Illustrated Flora of the Northeastern United States and Adjacent Canada. Vol. 2. Hafner, New York. 655 p., p. 188.
  2. ^ "Podophyllum peltatum". Integrated Taxonomic Information System. 
  3. ^ a b Fondren, Brian T. "Mayapple". Ethnobotanical leaflets. Retrieved 2006-06-03. 
  4. ^ a b Keddy, P.A. 2007. Plants and Vegetation: Origins, Processes, Consequences. Cambridge University Press, Cambridge, UK. 666 p.
  5. ^ Sohn, J.J. and D. Policansky. 1977. The Costs of Reproduction in the Mayapple Podophyllum Peltatum (Berberidaceae). Ecology 58:1366–1374. http://dx.doi.org/10.2307/1935088
  6. ^ Watson, M.A. and five others. 2001. The developmental ecology of mycorrhizal associations in mayapple, Podophyllum peltatum, Berberidaceae. Evolutionary Ecology 15: 425-442.
  7. ^ "Puccinia podophylli Schwein. Mayapple rust". Iowa State University, Ada Hayden Herbarium (ISC). Retrieved 2013-03-16. 
  8. ^ Bunyard, Britt A. 2013 "Mayapple Rust Resurrection" FUNGI 6(1): 38-39.
  9. ^ Podophyllum peltatum at USDA PLANTS Database
  10. ^ Blanchan, Neltje (2002). Wild Flowers: An Aid to Knowledge of our Wild Flowers and their Insect Visitors. Project Gutenberg Literary Archive Foundation. 
  11. ^ Moraes, R.M., H. Lata, E. Bedir, M. Maqbool, and K. Cushman. 2002. On American Mayapple as practical source of podophyllotoxin p. 527–532. In: J. Janick and A. Whipkey (eds.), Trends in new crops and new uses. ASHS Press, Alexandria, VA.
  12. ^ Brunton LL et al. Goodman and Gilman's The Pharmacological Basis of Therapeutics, chapter: 61. Cytotoxic agents/Epipodophyllotoxins Twelfth Edition ISBN 978-0-07-162442-8
  13. ^ Lewis, W.H. and M.P.F. Elvin-Lewis. 1977. Medical Botany. Plants Affecting Man's Health. Wiley, New York. 515 p. p. 123-124.
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Notes

Comments

The following forms have been described: 

 Podophyllum peltatum forma aphyllum Plitt--fertile shoots with no foliage leaves; Podophyllum peltatum forma biltmoreanum Steyermark--fruits orange; Podophyllum peltatum forma deamii Raymond--fruits and seeds maroon, and flowers, placentae, and plant axes pink-tinged; Podophyllum peltatum forma polycarpum (Clute) Plitt--flowers with multiple, free carpels.

The ripe fruit of Podophyllum peltatum is considered edible; all other parts of the plant are toxic. Several lignans and their glycosides, present in the resin extracted from rhizomes and roots, exhibit antitumor activity. Etoposide, a semisynthetic derivative of one of the lignans, is currently used in the treatment of small-cell lung cancer and testicular cancer (P. M. Dewick 1983). Native Americans used Podophyllum for a wide variety of medicinal purposes and as an insecticide (D. E. Moerman 1986).

Podophyllum peltatum is sometimes cultivated in woodland gardens, and some populations on the periphery of its geographical range may be escapes from cultivation.

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