''Maianthemum'' includes the former genus ''Smilacina'' and is a

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...

herbaceous Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition of t ...

flowering plants with fleshy, persistent

rhizome In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow hori ...

s. It is widespread across much of

North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...

Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...

and

Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...

,Kew World Checklist of Selected Plant Families
/ref>Flora of North America, Vol. 26 Page 206, ''Maianthemum'' F. H. Wiggers, Prim. Fl. Holsat. 14. 1780.
/ref> and may be terrestrial, aquatic or

epiphytic An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...

. It is characterized by simple, unbranched stems that are upright, leaning or hanging down and have 2–17 foliage leaves. Leaves are simple and may clasp the stem or be short-

petiolate Petiole may refer to: *Petiole (botany), the stalk of a leaf, attaching the blade to the stem *Petiole (insect anatomy) In entomology, petiole is the technical term for the narrow waist of some hymenopteran insects, especially ants, bees, a ...

. The

inflorescence An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...

is terminal and either a

panicle A panicle is a much-branched inflorescence. (softcover ). Some authors distinguish it from a compound spike inflorescence, by requiring that the flowers (and fruit) be pedicellate (having a single stem per flower). The branches of a panicle are of ...

or a

raceme A raceme ( or ) or racemoid is an unbranched, indeterminate type of inflorescence bearing flowers having short floral stalks along the shoots that bear the flowers. The oldest flowers grow close to the base and new flowers are produced as the s ...

with few to many

pedicel Pedicle or pedicel may refer to: Human anatomy *Pedicle of vertebral arch, the segment between the transverse process and the vertebral body, and is often used as a radiographic marker and entry point in vertebroplasty and kyphoplasty procedures ...

ate flowers. Most

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

have 6

tepals A tepal is one of the outer parts of a flower (collectively the perianth). The term is used when these parts cannot easily be classified as either sepals or petals. This may be because the parts of the perianth are undifferentiated (i.e. of very ...

and 6

stamens The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...

; a few have parts in 4s. Tepals are distinct in most species and all of similar size. Flowers are spreading, cup-shaped or bell-shaped and usually white, but lavender to red or green in some species. Fruits are rounded to lobed berries containing few to several seeds.

Vegetative

; Rhizomes and roots Rhizomes are persistent and have scale leaves. They can be

sympodial Sympodial growth is a bifurcating branching pattern where one branch develops more strongly than the other, resulting in the stronger branches forming the primary shoot and the weaker branches appearing laterally. A sympodium, also referred to a ...

; spreading and thread-like, or cylindric, clumped and fleshy. ''

Maianthemum paludicola ''Maianthemum paludicola'' is a perennial flowering plant. It is a rare terrestrial herb, endemic to Costa Rica. It has only been found in high-elevation bogs and wetlands and was first described in 1986. Description Flower- and leaf-bearing stem ...

'' has an unusual woody, upright sympodial rhizome set above ground. Roots may be spread along the rhizome, clumped at the nodes, or clumped near the base of leafy shoots. The rhizome is the perennial part of the plant and growth is by branching of the rhizome. Flowering/fruiting shoots are attached to the rhizome by a discreet internode, are short-lived and wither at the end of the growing season. ; Stems and leaves The stems are unbranched and pendant to arching or upright. Fertile plants have 2 to 17 or more stem (cauline)

leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...

(arising from the stem) that are alternate along the stem. Most species produce fertile shoots seasonally. A few species such as ''

Maianthemum canadense ''Maianthemum canadense'' (Canadian may-lily, Canada mayflower, false lily-of-the-valley, Canadian lily-of-the-valley, wild lily-of-the-valley,, p.105 two-leaved Solomon's seal) is an understory perennial flowering plant, native to Canada and the ...

'' and ''

Maianthemum dilatatum ''Maianthemum dilatatum'' (snakeberry, two-leaved Solomon's seal or false lily of the valley) is a common rhizomatous perennial flowering plant that is native to western North America from northern California to the Aleutian islands, and Asia acr ...

'' also have a vegetative phase, producing a solitary leaf on sterile shoots; plants may continue in this vegetative phase for up to six years. ''

Maianthemum trifolium ''Maianthemum trifolium'' (Three-leaf Solomon's-seal, three-leaf Solomon's-plume, threeleaf false lily of the valley, smilacine trifoliée) is a species of flowering plant that is associated with extremely wet environments and is native to Canada ...

'' produces two petiolate foliage leaves the first year, then a flowering shoot the second year with 2-4 sessile leaves. In most species, leaves have a large midvein, dividing the leaves in half, then each half is again divided by a slightly smaller vein; each quarter again divided by progressively finer veins. The resulting pattern is that the leaves have three prominent veins, and adjacent veins are of different diameters. A few species, such as ''

Maianthemum stellatum ''Maianthemum stellatum'' (star-flowered, starry, or little false Solomon's seal, or simply false Solomon's seal; star-flowered lily-of-the-valley or starry false lily of the valley; syn. ''Smilacina stellata'') is a species of flowering plant, ...

'', have leaves with veins that are often all of roughly the same diameter. Leaf edges may be flat or wavy (undulate) and sometimes have small, outward-pointing teeth.

Reproductive

; Inflorescence The

is either a

or a

with few to many

late flowers. Racemes may be simple (one flower per node) or complex (some nodes with 2 or more flowers). Panicles may be few- to many-branched and the branches themselves are often racemose. A few Asian species such as ''

Maianthemum henryi ''Maianthemum'' includes the former genus ''Smilacina'' and is a genus of perennial herbaceous flowering plants with fleshy, persistent rhizomes. It is widespread across much of North America, Europe and Asia, and may be terrestrial, aquatic or E ...

'' have a main axis with fully extended racemose branches at lower nodes and upper nodes with clustered flowers. The type and pattern of inflorescence is the most useful feature to separate ''Maianthemum'' species. ; Flowers The flowers are

actinomorphic Floral symmetry describes whether, and how, a flower, in particular its perianth, can be divided into two or more identical or mirror-image parts. Uncommonly, flowers may have no axis of symmetry at all, typically because their parts are spirall ...

(radially symmetric) and relatively small. The

perianth The perianth (perigonium, perigon or perigone in monocots) is the non-reproductive part of the flower, and structure that forms an envelope surrounding the sexual organs, consisting of the calyx (sepals) and the corolla (petals) or tepals when ...

in most species has six

, in two whorls of three (

trimerous Merosity (from the greek "méros," which means "having parts") refers to the number of component parts in a distinct whorl of a plant structure. The term is most commonly used in the context of a flower where it refers to the number of sepals in a ...

). ''

'', and ''

Maianthemum bifolium ''Maianthemum bifolium'' (false lily of the valley or May lily) is often a localized common rhizomatous flowering plant, native from western Europe (including Britain''New Flora of The British Isles'', Clive Stace) east to Siberia, China and Japa ...

'' however are

dimerous Merosity (from the greek "méros," which means "having parts") refers to the number of component parts in a distinct whorl of a plant structure. The term is most commonly used in the context of a flower where it refers to the number of sepals in a w ...

, with 4-tepaled flowers. Tepals are greenish, or white to pink or mauve in most species, but are a deep purple in '' Maianthemum purpureum'' and may have purple spots in '' Maianthemum gigas var. gigas''. The tepals in most species are free (not fused), all of a similar size and flowers are spreading to cup-shaped. Some Asian species such as ''Maianthemum henryi'' have bell-shaped (campanulate) flowers with partially fused tepals. The flowers are usually

bisexual Bisexuality is a romantic or sexual attraction or behavior toward both males and females, or to more than one gender. It may also be defined to include romantic or sexual attraction to people regardless of their sex or gender identity, whi ...

, containing both male (

androecium The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...

) and female (

gynoecium Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils'' ...

) parts, although two species from Japan, '' Maianthemum hondoense'' and '' Maianthemum yesoense'', are reported to be

unisexual Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...

. ; Gynoecium The

ovaries The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. T ...

are superior (attached above the other floral parts) and are spherical to cylindrical. The

pistil Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils'' ...

has two or three carpels separated by walls with nectar-carrying canals.

Style Style is a manner of doing or presenting things and may refer to: * Architectural style, the features that make a building or structure historically identifiable * Design, the process of creating something * Fashion, a prevailing mode of clothing ...

s are narrow (< 5mm wide) and about as long as the ovary. Stigmas are lobed and slightly wider than the style. ; Androecium

Stamens The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...

are 4 or 6 and usually inserted at the base of the tepals, but may be inserted about 1 mm above, as in '' Maianthemum flexuosum''. Anthers are cylindrical and split longitudinally. ; Fruit and seeds The

berry A berry is a small, pulpy, and often edible fruit. Typically, berries are juicy, rounded, brightly colored, sweet, sour or tart, and do not have a stone or pit, although many pips or seeds may be present. Common examples are strawberries, raspb ...

is globose and often lobed. Berries ripen to red. In most species unripe berries are green or green mottled with red or copper. In some, e.g. ''

'' the immature berries have dark stipes. The

seeds A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosperm pl ...

are usually few and spherical but in some species may be conical.

Morphology

file:Maianthemum stellatum rhizomes.jpg, Rhizome and roots: ''M. stellatum'' file:Maianthemum racemosum amplexicaule blooming plant.jpg, Unbranched, upright stem: ''M. racemosum amplexicaule'' file:Maianthemum canadense sterile plants.jpg, Sterile plants: ''M. canadense'' file:Maianthemum stellatum 15290.JPG, Simple raceme; one pedicellate flower per node: ''M. stellatum'' file:Flower buds close up Maianthemum canadense.jpg, Complex raceme, some nodes with 2 flowers: ''M. canadense'' file:Maian amplex panicle.jpg, Panicle with main axis and side branches: ''M. racemosum amplexicaule''

Taxonomy

The current concept includes species that were previously divided into 2 closely related genera; ''Maianthemum'', including dimerous species and ''Smilacina'' for the trimerous ones. Studies in the 1970s showed strong genetic similarity, similar fruits, and evidence that the 4-tepal species evolved from a 6-tepal species, so the genus ''Smilacina'' was combined with ''Maianthemum''.

Phylogeny

In the APG III classification system, ''Maianthemum'' is placed in the

family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...

Asparagaceae Asparagaceae, known as the asparagus family, is a family of flowering plants, placed in the order Asparagales of the monocots. The family name is based on the edible garden asparagus, ''Asparagus officinalis''. Those who live in the temperate c ...

, subfamily

Nolinoideae Nolinoideae is a monocot subfamily of the family (biology), family Asparagaceae in the APG III system of 2009. It used to be treated as a separate family, Ruscaceae sensu lato, s.l. The family name is derived from the Binomial nomenclature, gener ...

(formerly the family Ruscaceae). Like many

lilioid monocot Lilioid monocots (lilioids, liliid monocots, petaloid monocots, petaloid lilioid monocots) is an informal name used for a grade (grouping of taxa with common characteristics) of five monocot orders (Petrosaviales, Dioscoreales, Pandanales, Lili ...

s, ''Maianthemum'' (including the former ''Smilacina'') were formerly included in the family

Liliaceae The lily family, Liliaceae, consists of about 15 genera and 610 species of flowering plants within the order Liliales. They are monocotyledonous, perennial, herbaceous, often bulbous geophytes. Plants in this family have evolved with a fair ...

. The genus has also been placed in the former family

Convallariaceae Nolinoideae is a monocot subfamily of the family Asparagaceae in the APG III system of 2009. It used to be treated as a separate family, Ruscaceae s.l. The family name is derived from the generic name of the type genus, ''Nolina''. The subfami ...

, and resembles the closely related ''

Polygonatum ''Polygonatum'' , also known as King Solomon's-seal or Solomon's seal, is a genus of flowering plants. In the APG III classification system, it is placed in the family Asparagaceae, subfamily Nolinoideae (formerly the family Ruscaceae). It has ...

'', hence the common name "false Solomon's seal".

Etymology

''Maianthemum'' comes from the Greek for "May" (maios) and “flower” (Anthos).

Distribution and habitat

The genus is widespread across much of

and

,Kew World Checklist of Selected Plant Families
/ref>Flora of North America, Vol. 26 Page 206, ''Maianthemum'' F. H. Wiggers, Prim. Fl. Holsat. 14. 1780.
/ref>

Ecology

The majority of species are spring-flowering forest herbs, requiring shaded, moist conditions and cool temperatures. Some however (e.g. '' Maianthemum monteverdense'') are epiphytes. Others such as ''

'' grow in diverse habitats from sand dunes to forest under-stories. And ''

'' and ''

'' are found in full sun in open wetlands. ''M. trifolium'' is sometimes considered aquatic. The species that are forest herbs generally begin growth in early spring, with leaves expanding before the tree canopy fully develops. Epiphytic tropical species usually grow new shoots in the dry season.

Conservation

Many of the North American ''Maianthemum'' are widespread and relatively common. Several though are considered endemic to the wider “Mesomexico”. ''Maianthemum comaltepecense'' is a rare perennial, terrestrial herb found as an understory species in moist forests, known only from Oaxaca, Mexico.Espejo et al.1996. Maianthemum comaltepecense, Una Nueva Especie del Estado de Oaxaca. Acta Botánica Mexicana (1996), 36:21-28 and

endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...

to southwest Mexico.

Species

Species accepted: #'' Maianthemum amoenum'' (H.L.Wendl.) LaFrankie – Honduras, El Salvador, Guatemala, Mexico (Chiapas, Oaxaca, Veracruz) # '' Maianthemum atropurpureum'' (Franch.) LaFrankie – Sichuan, Yunnan #'' Maianthemum bicolor'' (Nakai) Cubey – Korea # ''

'' (L.) F.W.Schmidt – northern and central Europe and northern Asia from Spain to Kamchatka; China, Japan, Siberia, Kazakhstan, Germany, Italy, Ukraine, etc # ''

'' Desf. – St. Pierre and Miquelon, much of eastern USA and all provinces and territories in Canada except Nunavut #'' Maianthemum comaltepecense'' Espejo, López-Ferr. & Ceja – Mexico (Oaxaca) # '' Maianthemum dahuricum'' (Turcz. ex Fisch. & C.A.Mey.) LaFrankie – Siberia, Russian Far East, Manchuria, Inner Mongolia, Korea # ''

'' (Alph.Wood) A.Nelson & J.F.Macbr. – Mongolia, Korea, Japan, Russian Far East, Alaska, Yukon, British Columbia, Washington, Idaho, Oregon, California #'' Maianthemum flexuosum'' (Bertol.) LaFrankie – Mexico (Chiapas, Guerrero, Hidalgo, Michoacán, Oaxaca, Veracruz), El Salvador, Guatemala, Honduras, Nicaragua #'' Maianthemum formosanum'' (Hayata) LaFrankie – Taiwan # '' Maianthemum forrestii'' (W.W.Sm.) LaFrankie – Yunnan # '' Maianthemum fusciduliflorum'' (Kawano) S.C.Chen & Kawano – Tibet, Yunnan, Bhutan, Assam, Myanmar # '' Maianthemum fuscum'' (Wall.) LaFrankie – Nepal, Tibet, Yunnan, Bhutan, Assam, Myanmar #'' Maianthemum gigas'' (Woodson) LaFrankie – Mexico (Chiapas), Costa Rica, Guatemala, Panama, and Nicaragua # '' Maianthemum gongshanense'' (S.Yun Liang) H.Li – Yunnan # ''

'' (Baker) LaFrankie – Vietnam, Myanmar, Tibet, southern China #'' Maianthemum hondoense'' (Ohwi) LaFrankie – Honshu #'' Maianthemum × intermedium'' Vorosch. – Siberia, Russian Far East ''(M. bifolium × M. dilatatum)'' # '' Maianthemum japonicum'' (A.Gray) LaFrankie – Russian Far East, Japan, Korea, northeastern China # '' Maianthemum lichiangense'' (W.W.Sm.) LaFrankie – Gansu, Shaanxi, Sichuan, Yunnan #'' Maianthemum macrophyllum'' (M.Martens & Galeotti) LaFrankie – Mexico (Veracruz, Oaxaca) #'' Maianthemum mexicanum'' García Arév – Mexico (Durango, Sinaloa) #'' Maianthemum monteverdense'' LaFrankie – Nicaragua, Costa Rica # '' Maianthemum nanchuanense'' H.Li & J.L.Huang – Sichuan # '' Maianthemum oleraceum'' (Baker) LaFrankie – Nepal, Bhutan, Assam, Myanmar, Tibet, Sikkim, Guizhou, Sichuan, Yunnan #''

'' LaFrankie – Costa Rica # '' Maianthemum paniculatum'' (M.Martens & Galeotti) LaFrankie – Mexico (Chiapas, Guerrero, Hidalgo, Oaxaca, Puebla and Veracruz), Costa Rica, El Salvador, Guatemala, Honduras, and Panamá # '' Maianthemum purpureum'' (Wall.) LaFrankie – Tibet, Yunnan, Nepal, Bhutan, Sikkim, Assam (also called Himalayan Mayflower or Purple Mayflower) # ''

Maianthemum racemosum ''Maianthemum racemosum'', the treacleberry, feathery false lily of the valley, false Solomon's seal, Solomon's plume or false spikenard, is a species of flowering plant native to North America. It is a common, widespread plant with numerous comm ...

'' (L.) Link – All USA states, all Canadian provinces and territories (except Nunavut and Labrador) and Mexico (Chihuahua) #'' Maianthemum robustum'' (Makino & Honda) LaFrankie – Honshu #'' Maianthemum salvinii'' (Baker) LaFrankie – Mexico (Chiapas), Guatemala #'' Maianthemum scilloideum'' (M.Martens & Galeotti) LaFrankie – Mexico (Chiapas, Coahuila, Guerrero, Michoacán, Nuevo León, Oaxaca, Puebla and Veracruz), Guatemala, Honduras # ''

'' (L.) Link – Most states in USA (except those southeast of New Mexico), all Canadian provinces and territories (except Nunavut and Labrador) and in the Mexican states of Chihuahua, Coahuila, Nuevo León and Sonoroa; introduced in Norway, Sweden # '' Maianthemum stenolobum'' (Franch.) S.C.Chen & Kawano – Sichuan, Gansu, Hubei # '' Maianthemum szechuanicum'' (F.T.Wang & Tang) H.Li – Sichuan, Yunnan # '' Maianthemum tatsienense'' (Franch.) LaFrankie – Bhutan, Assam, Myanmar, Gansu, Guangxi, Guizhou, Hubei, Hunan, Sichuan, Yunnan # ''

'' (L.) Sloboda – Siberia, Russian Far East to North Korea, St. Pierre and Miquelon, all Canadian provinces and territories, northeastern United States # '' Maianthemum tubiferum'' (Batalin) LaFrankie – Gansu, Hubei, Qinghai, Shaanxi, Sichuan #'' Maianthemum yesoense'' (Franch. & Sav.) LaFrankie – Japan

References

* {{Taxonbar, from=Q157848 Asparagaceae genera