Alder
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Alders are trees of the genus Alnus in the birch family Betulaceae. The genus includes about 35 species[1] of monoecious trees and shrubs, a few reaching a large size, distributed throughout the north temperate zone with a few species extending into Central America, as well as the northern and southern Andes.[2]
Description
With a few exceptions, alders are deciduous, and the leaves are alternate, simple, and serrated. The flowers are catkins with elongate male catkins on the same plant as shorter female catkins, often before leaves appear; they are mainly wind-pollinated, but also visited by bees to a small extent. These trees differ from the birches (Betula, another genus in the family) in that the female catkins are woody and do not disintegrate at maturity, opening to release the seeds in a similar manner to many conifer cones.
The largest species are red alder (A. rubra) on the west coast of North America, and black alder (A. glutinosa), native to most of Europe and widely introduced elsewhere, both reaching over Template:Convert. By contrast, the widespread Alnus alnobetula (green alder) is rarely more than a Template:Convert shrub.
Phylogeny
Classification
The genus is divided into three subgenera:
Subgenus Alnus
Trees with stalked shoot buds, male and female catkins produced in autumn (fall) but stay closed over winter, pollinating in late winter or early spring, about 15–25 species, including:
- Alnus acuminata KunthScript error: No such module "Check for unknown parameters".
- subsp. acuminata KunthScript error: No such module "Check for unknown parameters".
- subsp. arguta (Schltdl.) FurlowScript error: No such module "Check for unknown parameters".
- subsp. glabrata (Fernald) FurlowScript error: No such module "Check for unknown parameters".
- Alnus cordata (Loisel.) DubyScript error: No such module "Check for unknown parameters".
- Alnus cremastogyne BurkillScript error: No such module "Check for unknown parameters".
- Alnus firma Siebold & Zucc.Script error: No such module "Check for unknown parameters".
- Alnus glutinosa (L.) Gaertn.Script error: No such module "Check for unknown parameters".
- subsp. barbata (C.A.Mey.) Yalt.Script error: No such module "Check for unknown parameters".
- subsp. glutinosa (L.) Gaertn.Script error: No such module "Check for unknown parameters".
- subsp. incisa (Willd.) RegelScript error: No such module "Check for unknown parameters".
- subsp. laciniata (Willd.) RegelScript error: No such module "Check for unknown parameters".
- Alnus hirsuta (Spach) Rupr.Script error: No such module "Check for unknown parameters".
- Alnus incana (L.) MoenchScript error: No such module "Check for unknown parameters".
- subsp. incana (L.) MoenchScript error: No such module "Check for unknown parameters".
- subsp. kolaensis (Orlova) Á.Löve & D.LöveScript error: No such module "Check for unknown parameters".
- subsp. rugosa (Du Roi) R.T.ClausenScript error: No such module "Check for unknown parameters".
- subsp. tenuifolia (Nutt.) BreitungScript error: No such module "Check for unknown parameters".
- Alnus japonica (Thunb.) Steud.Script error: No such module "Check for unknown parameters".
- Alnus jorullensis KunthScript error: No such module "Check for unknown parameters".
- subsp. lutea FurlowScript error: No such module "Check for unknown parameters".
- subsp. jorullensis KunthScript error: No such module "Check for unknown parameters".
- Alnus lusitanica Vít, Douda, & MandákScript error: No such module "Check for unknown parameters".[3]
- Alnus matsumurae CallierScript error: No such module "Check for unknown parameters".
- Alnus nepalensis D.DonScript error: No such module "Check for unknown parameters".
- Alnus oblongifolia Torr.Script error: No such module "Check for unknown parameters".
- Alnus orientalis Decne.Script error: No such module "Check for unknown parameters".
- Alnus rhombifolia Nutt.Script error: No such module "Check for unknown parameters".
- Alnus rohlenae Vít, Douda, & MandákScript error: No such module "Check for unknown parameters".[3]
- Alnus rubra Bong.Script error: No such module "Check for unknown parameters".
- Alnus serrulata (Aiton) Willd.Script error: No such module "Check for unknown parameters".
- Alnus subcordata C.A.Mey.Script error: No such module "Check for unknown parameters".
- Alnus tenuifolia Nutt.Script error: No such module "Check for unknown parameters".
- Alnus trabeculosa Hand.-Mazz.Script error: No such module "Check for unknown parameters".
Subgenus Clethropsis
Trees or shrubs with stalked shoot buds, male and female catkins produced in autumn (fall) and expanding and pollinating then, three species:
- Alnus formosana (Burkill) MakinoScript error: No such module "Check for unknown parameters".
- Alnus maritima (Marshall) Muhl. ex Nutt.Script error: No such module "Check for unknown parameters".
- Alnus nitida (Spach) Endl.Script error: No such module "Check for unknown parameters".
Subgenus Alnobetula
Shrubs with shoot buds not stalked, male and female catkins produced in late spring (after leaves appear) and expanding and pollinating then, one to four species:
- Alnus alnobetula (Ehrh.) K.KochScript error: No such module "Check for unknown parameters". (synonym-Alnus viridis)
- subsp. alnobetula (Ehrh.) K.KochScript error: No such module "Check for unknown parameters".
- subsp. crispa (Aiton) RausScript error: No such module "Check for unknown parameters".
- subsp. fruticosa (Rupr.) RausScript error: No such module "Check for unknown parameters".
- subsp. sinuata (Regel) RausScript error: No such module "Check for unknown parameters".
- subsp. suaveolens (Req.) Lambinon & KerguélenScript error: No such module "Check for unknown parameters".
- Alnus firma Siebold & Zucc.Script error: No such module "Check for unknown parameters".
- Alnus mandshurica (Callier) Hand.-Mazz.Script error: No such module "Check for unknown parameters".
- Alnus maximowiczii CallierScript error: No such module "Check for unknown parameters".
- Alnus pendula Matsum.Script error: No such module "Check for unknown parameters".
- Alnus sieboldiana Matsum.Script error: No such module "Check for unknown parameters".
Not assigned to a subgenus
- Alnus fauriei H.Lév. & VaniotScript error: No such module "Check for unknown parameters".
- Alnus ferdinandi-coburgii C.K.Schneid.Script error: No such module "Check for unknown parameters".
- Alnus glutipes (Jarm. ex Czerpek) Vorosch.Script error: No such module "Check for unknown parameters".
- Alnus hakkodensis HayashiScript error: No such module "Check for unknown parameters".
- Alnus henryi C.K.Schneid.Script error: No such module "Check for unknown parameters".
- Alnus lanata Duthie ex BeanScript error: No such module "Check for unknown parameters".
- Alnus mairei H.Lév.Script error: No such module "Check for unknown parameters".
- Alnus paniculata NakaiScript error: No such module "Check for unknown parameters".
- Alnus serrulatoides CallierScript error: No such module "Check for unknown parameters".
- Alnus vermicularis NakaiScript error: No such module "Check for unknown parameters".
Species names with uncertain taxonomic status
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- Alnus balatonialis BorbásScript error: No such module "Check for unknown parameters".
- Alnus cuneata Geyer ex Walp.Script error: No such module "Check for unknown parameters".
- Alnus dimitrovii Jordanov & KitanovScript error: No such module "Check for unknown parameters".
- Alnus djavanshirii H.ZareScript error: No such module "Check for unknown parameters". – Iran
- Alnus dolichocarpa H.Zare, Amini & AssadiScript error: No such module "Check for unknown parameters". – Iran
- Alnus figerti CallierScript error: No such module "Check for unknown parameters".
- Alnus frangula L. ex HuthScript error: No such module "Check for unknown parameters".
- Alnus gigantea NakaiScript error: No such module "Check for unknown parameters".
- Alnus glandulosa Sarg.Script error: No such module "Check for unknown parameters".
- Alnus henedae Sugim.Script error: No such module "Check for unknown parameters".
- Alnus hybrida Rchb.Script error: No such module "Check for unknown parameters".
- Alnus laciniata Ehrh.Script error: No such module "Check for unknown parameters".
- Alnus lobata NymanScript error: No such module "Check for unknown parameters".
- Alnus microphylla Arv.-Touv.Script error: No such module "Check for unknown parameters".
- Alnus obtusifolia Mert. ex RegelScript error: No such module "Check for unknown parameters".
- Alnus oxyacantha LavalleScript error: No such module "Check for unknown parameters".
- Alnus subrotunda Desf.Script error: No such module "Check for unknown parameters".
- Alnus vilmoriana LebasScript error: No such module "Check for unknown parameters".
- Alnus washingtonia WetzelScript error: No such module "Check for unknown parameters".
Hybrids
The following hybrids have been described:[4][5]
- Alnus × elliptica Req.Script error: No such module "Check for unknown parameters". (A. cordata × A. glutinosa)
- Alnus × fallacina CallierScript error: No such module "Check for unknown parameters". (A. incana subsp. rugosa × A. serrulata)
- Alnus × hanedae SuyinataScript error: No such module "Check for unknown parameters". (A. firma × A. sieboldiana)
- Alnus × hosoii Mizush.Script error: No such module "Check for unknown parameters". (A. maximowiczii × A. pendula)
- Alnus × mayrii CallierScript error: No such module "Check for unknown parameters". (A. hirsuta × A. japonica)
- Alnus × peculiaris HiyamaScript error: No such module "Check for unknown parameters". (A. firma × A. pendula)
- Alnus × pubescens Tausch.Script error: No such module "Check for unknown parameters". (A. glutinosa × A. incana)
- Alnus × suginoi Sugim.Script error: No such module "Check for unknown parameters".
The status of the following hybrids is unresolved:[4]
- Alnus × aschersoniana CallierScript error: No such module "Check for unknown parameters".
- Alnus × koehnei CallierScript error: No such module "Check for unknown parameters".
- Alnus × ljungeri MuraiScript error: No such module "Check for unknown parameters".
- Alnus × purpusii CallierScript error: No such module "Check for unknown parameters".
- Alnus × silesiaca FiekScript error: No such module "Check for unknown parameters".
- Alnus × spaethii CallierScript error: No such module "Check for unknown parameters". (A. japonica × A. subcordata)
Fossil record
The oldest fossil pollen that can be identified as Alnus is from northern Bohemia, dating to the late Paleocene, around 58 million years ago.[6]
- †Alnus fairi (Knowlton) Wolfe, 1966Script error: No such module "Check for unknown parameters". - Miocene; Western North America[7]
- †Alnus heterodonta (Newberry) Meyer & Manchester 1987Script error: No such module "Check for unknown parameters". – Oligocene; Fossil, Oregon
- †Alnus hollandiana Jennings, 1920Script error: No such module "Check for unknown parameters". - Miocene; Western North America[8]
- †Alnus largei (Knowlton) Wolfe, 1966Script error: No such module "Check for unknown parameters". - Miocene; Western North America[7]
- †Alnus parvifolia (Berry) Wolfe & Wehr, 1987Script error: No such module "Check for unknown parameters". - Ypresian; Okanagan Highlands[9]
- †Alnus relatus (Knowlton) Brown, 1937Script error: No such module "Check for unknown parameters". - Miocene; Western North America[7]
Etymology
The common name alder evolved from the Old English word alor, which in turn is derived from Proto-Germanic root aliso.[10] The generic name Alnus is the equivalent Latin name, from whence French aulne and Spanish Alamo (Spanish term for "poplar").[10]
Ecology
Alders are commonly found near streams, rivers, and wetlands. Sometimes where the prevalence of alders is particularly prominent these are called alder carrs. In the Pacific Northwest of North America, the white alder (Alnus rhombifolia) unlike other northwest alders, has an affinity for warm, dry climates, where it grows along watercourses, such as along the lower Columbia River east of the Cascades and the Snake River, including Hells Canyon.
Alder leaves and sometimes catkins are used as food by numerous butterflies and moths.
A. glutinosa and A. viridis are classed as environmental weeds in New Zealand.[11] Alder leaves and especially the roots are important to the ecosystem because they enrich the soil with nitrogen and other nutrients.
Nitrogen fixation and succession of woodland species
Alder is particularly noted for its important symbiotic relationship with Frankia alni, an actinomycete, filamentous, nitrogen-fixing bacterium. This bacterium is found in root nodules, which may be as large as a human fist, with many small lobes, and light brown in colour. The bacterium absorbs nitrogen from the air and makes it available to the tree. Alder, in turn, provides the bacterium with sugars, which it produces through photosynthesis. As a result of this mutually beneficial relationship, alder improves the fertility of the soil where it grows, and as a pioneer species, it helps provide additional nitrogen for the successional species to follow.
Because of its abundance, red alder delivers large amounts of nitrogen to enrich forest soils. Red alder stands have been found to supply between Template:Convert of nitrogen annually to the soil. From Alaska to Oregon, Alnus viridis subsp. sinuata (A. sinuata, Sitka alder or slide alder), characteristically pioneer fresh, gravelly sites at the foot of retreating glaciers. Studies show that Sitka alder, a more shrubby variety of alder, adds nitrogen to the soil at an average rate of Template:Convert per year, helping convert the sterile glacial terrain to soil capable of supporting a conifer forest. Alders are common among the first species to colonize disturbed areas from floods, windstorms, fires, landslides, etc. Alder groves often serve as natural firebreaks since these broad-leaved trees are much less flammable than conifers. Their foliage and leaf litter does not carry a fire well, and their thin bark is sufficiently resistant to protect them from light surface fires. In addition, the light weight of alder seedsTemplate:Sndnumbering Template:ConvertTemplate:Sndallows for easy dispersal by the wind. Although it outgrows coastal Douglas-fir for the first 25 years, it is very shade intolerant and seldom lives more than 100 years. Red alder is the Pacific Northwest's largest alder and the most plentiful and commercially important broad-leaved tree in the coastal Northwest. Groves of red alder Template:Convert in diameter intermingle with young Douglas-fir forests west of the Cascades, attaining a maximum height of Template:Convert in about sixty years, and then are afflicted by heart rot. Alders largely help create conditions favorable for giant conifers that replace them.[1]
- Alder root nodules
-
Whole root nodule
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Sectioned root nodules
Parasites
Alder roots are parasitized by northern groundcone.
Uses
The catkins of some alder species have a degree of edibility,[12] and may be rich in protein. Reported to have a bitter and unpleasant taste, they are more useful for survival purposes. The wood of certain alder species is often used to smoke various food items such as coffee, salmon, and other seafood.
Alder is notably stable when immersed, and has been used for millennia as a material for pilings for piers and wharves. Most of the pilings that form the foundation of Venice were made from alder trees.[13]
Alder bark contains the anti-inflammatory salicin, which is metabolized into salicylic acid in the body.[14] Some Native American cultures use red alder bark (Alnus rubra) to treat poison oak, insect bites, and skin irritations. Blackfeet Indians have traditionally used an infusion made from the bark of red alder to treat lymphatic disorders and tuberculosis. Recent clinical studies have verified that red alder contains betulin and lupeol, compounds shown to be effective against a variety of tumors.[15]
The inner bark of the alder, as well as red osier dogwood, or chokecherry, is used by some Indigenous peoples of the Americas in smoking mixtures, known as kinnikinnick, to improve the taste of the bearberry leaf.[16]
Alder is illustrated in the coat of arms for the Austrian town of Grossarl.
Electric guitars, most notably those manufactured by the Fender Musical Instruments Corporation, have been built with alder bodies since the 1950s. Alder is appreciated for its tone that is claimed to be tight and evenly balanced, especially when compared to mahogany, and has been adopted by many electric guitar manufacturers. It usually is finished in opaque lacquer (nitrocellulose, polyurethane, or polyester), as it does not have a prominent grain.
As a hardwood, alder is used in making furniture, cabinets, and other woodworking products. In these applications, its aforementioned lack of prominent grain means that it is often veneered, either by stained light woods such as oak, ash, or figured maple, or by darker woods such as teak or walnut.
Alder bark and wood (like oak and sweet chestnut) contain tannin and are traditionally used to tan leather.
A red dye can also be extracted from the outer bark, and a yellow dye from the inner bark.[17]
Culture
Ermanno Olmi's movie The Tree of Wooden Clogs (L' Albero Degli Zoccoli, 1978) refers in its title to alder, typically used to make clogs as in this movie's plot.[18][19]
References
Further reading
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External links
Template:Sister project Template:Sister project
- Flora Europaea: Alnus
- Flora of Bolivia: Alnus
- Flora of China: Alnus
- Flora of North America: Alnus
- Flora of Pakistan: Alnus
Template:Tannin source Template:Taxonbar Template:Authority control
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- ↑ PRESSO LA RIVA: L'ONTANO (tr. AT THE SHORE: THE ALDER), December 2015 www.officinadellambiente.com, accessed 17 November 2020
- ↑ Ontano nero (tr. Black Alder) accessed 17 November 2020 uomoenatura.it