Malus sylvestris
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''Malus sylvestris'', the European crab apple, also known as the European wild apple or simply the crab apple, is a
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of the genus ''
Malus ''Malus'' ( or ) is a genus of about 32–57 species of small deciduous trees or shrubs in the family Rosaceae, including the domesticated orchard apple, crab apples (sometimes known in North America as crabapples) and wild apples. The genus i ...
''. Its
scientific name In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
means "forest apple", reflecting its habitat. It is native to western Eurasia.


Description

The wild apple is a
deciduous In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed Leaf, leaves, usually in the autumn; to the shedding of petals, aft ...
small to medium-sized tree, but can also grow into a multi-stemmed bush. It can live 80–100 years and grow up to tall with trunk diameters of usually , although diameters exceeding have been recorded. The leaves are roundish-oval and sometimes hairy on the underside. The
hermaphrodite A hermaphrodite () is a sexually reproducing organism that produces both male and female gametes. Animal species in which individuals are either male or female are gonochoric, which is the opposite of hermaphroditic. The individuals of many ...
flowers appear in May, slightly preceding hawthorn, have white or pinkish
petal Petals are modified leaves that form an inner whorl surrounding the reproductive parts of flowers. They are often brightly coloured or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corol ...
s and are insect-pollinated. The small pome-fruits are around in diameter, ripen in autumn and fall to the ground. The bark is light brown and breaks up in flakes. The branches are spiny, especially in response to pruning or
browsing Browsing is a kind of orienting strategy. It is supposed to identify something of relevance for the browsing organism. In context of humans, it is a metaphor taken from the animal kingdom. It is used, for example, about people browsing open sh ...
.


Identification

European wild apples hybridise readily with domesticated apples, which can make identification difficult, as hybrids commonly exhibit transitional traits from both parent species. While certain identification relies on
genetic testing Genetic testing, also known as DNA testing, is used to identify changes in DNA sequence or chromosome structure. Genetic testing can also include measuring the results of genetic changes, such as RNA analysis as an output of gene expression, or ...
, field identification can be made by examining the following features. The crown of wild apples is densely branched, whereas the crown of domesticated apples tends to be more loosely arranged with fewer, straighter branches. Wild apple leaves tend to be somewhat smaller, stiffer and shinier. The underside of wild apple leaves also tends to be less hairy than those of both domesticated and hybrid apples, often lacking hairs altogether. Finally, wild apple fruits tend to be smaller, usually below in diameter, while domesticated and hybrid apples tend to be larger.


Taxonomy

Its
scientific name In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
means "forest apple".


Genetics and postglacial recolonization

Like most European tree species, the distribution of the European wild apple was limited to refugia in southern Europe during the Last Glacial Period. For the European wild apple, these refugia seem to have constituted southern France and northern Spain, the
Balkans The Balkans ( , ), corresponding partially with the Balkan Peninsula, is a geographical area in southeastern Europe with various geographical and historical definitions. The region takes its name from the Balkan Mountains that stretch throug ...
and possibly the
Carpathians The Carpathian Mountains or Carpathians () are a range of mountains forming an arc across Central Europe and Southeast Europe. Roughly long, it is the third-longest European mountain range after the Urals at and the Scandinavian Mountains ...
, respectively. From there, it recolonized the rest of Europe following the glacial retreat at the onset of the
Holocene The Holocene () is the current geologic time scale, geological epoch, beginning approximately 11,700 years ago. It follows the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene to ...
, and the colonization paths find themselves reflected in its modern distribution and genetic structure. Again as in many other organisms in Europe, plants and animals alike, the population of the European wild apple is divided into a large western population and a more strongly differentiated eastern population. Western Europe and northern Europe were most likely colonized from southern France, while eastern Europe was colonized from the Carpathians. In the process, some admixture between both populations seems to have occurred as they met.


Progenitor of cultivated apples

In the past ''M. sylvestris'' was thought to be the most important ancestor of the cultivated
apple An apple is a round, edible fruit produced by an apple tree (''Malus'' spp.). Fruit trees of the orchard or domestic apple (''Malus domestica''), the most widely grown in the genus, are agriculture, cultivated worldwide. The tree originated ...
(''M. domestica''), which has since been shown to have been primarily derived from the central Asian species '' M. sieversii''. As confirmed by DNA analyses, ''M. sylvestris'' has contributed significantly to the genome.Coart, E., Van Glabeke, S., De Loose, M., Larsen, A.S., Roldán-Ruiz, I. 2006. Chloroplast diversity in the genus ''Malus'': new insights into the relationship between the European wild apple (''Malus sylvestris'' (L.) Mill.) and the domesticated apple (''Malus domestica'' Borkh.). ''Mol. Ecol.'' 15 (8): 2171-82. Secondary
introgression Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Introg ...
from other species of the genus ''Malus'' has greatly shaped the genome of ''M. domestica'', with ''M. sylvestris'' being the largest secondary contributor. Current populations of ''M. domestica'' are more closely related to ''M. sylvestris'' than to ''M. sieversii'', while in more pure strains of ''M. domestica'', the ''M. sieversii'' ancestry still predominates.


Distribution and habitat

The tree is widespread throughout Europe, with the exception of the extreme north and south. Additionally, it is also distributed in
Anatolia Anatolia (), also known as Asia Minor, is a peninsula in West Asia that makes up the majority of the land area of Turkey. It is the westernmost protrusion of Asia and is geographically bounded by the Mediterranean Sea to the south, the Aegean ...
and further into the
south Caucasus The South Caucasus, also known as Transcaucasia or the Transcaucasus, is a geographical region on the border of Eastern Europe and West Asia, straddling the southern Caucasus Mountains. The South Caucasus roughly corresponds to modern Armenia, ...
. It occurs in a scattered distribution pattern as single individuals or in small groups. Due to its weak competitiveness and high light requirement, the wild apple is found mostly at sites where competition is reduced, such as the wet edge of forests, in wood pasture, farmland hedges or on very extreme, marginal sites. In the British Isles it occurs throughout
Ireland Ireland (, ; ; Ulster Scots dialect, Ulster-Scots: ) is an island in the North Atlantic Ocean, in Northwestern Europe. Geopolitically, the island is divided between the Republic of Ireland (officially Names of the Irish state, named Irelan ...
, England and Wales and with reduced frequency in Scotland.Worrel, Rick; Ruhsam, Markus; Renny, James; Jessop, Will; Findlay, Graeme (2018):
The Ecology and Genetics of Scotland's native wild apple: Malus sylvestris
In Scotland, it is most commonly found in woodlands and wood pastures.


Ecology and threats

The European wild apple is dispersed primarily by mammals, which eat the fruits. In a study from Mols, Denmark, it was determined that cattle accounted for the bulk of dispersal, followed by horses, despite the presence of wild animals. In Britain, the crab apple is associated with 93 species of insect. Its leaves are food of the hawthorn moth (''Scythropia crataegella''). Throughout its range, the wild apple is threatened and rare. Threats include the
introgression Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Introg ...
from domesticated apples, the lack of natural regeneration and modern forestry practice, which promotes the closure of forest canopy cover, as opposed to the formerly prevailing
coppice Coppicing is the traditional method in woodland management of cutting down a tree to a tree stump, stump, which in many species encourages new Shoot (botany), shoots to grow from the stump or roots, thus ultimately regrowing the tree. A forest ...
.Ulrike Hoffmann:
Wildäpfel im Spannungsfeld menschlichen Wirtschaftens
'. Natur in NRW 2/2018: 17-21. (In German)
In many aspects, the wild apple exhibits adaptations to
grazing In agriculture, grazing is a method of animal husbandry whereby domestic livestock are allowed outdoors to free range (roam around) and consume wild vegetations in order to feed conversion ratio, convert the otherwise indigestible (by human diges ...
and the presence of large herbivores, and consequently also a high degree of dependence on them. It is thorny, suitable for coppice due to pronounced resprouting abilities and a very hardy tree. Wild apples may survive crown collapse and the breakage of major branches. In dispersal, it appears to be strongly reliant on bovines and
equines ''Equus'' () is a genus of mammals in the perissodactyl family (biology), family Equidae, which includes wild horse, horses, Asinus, asses, and zebras. Within the Equidae, ''Equus'' is the only recognized Extant taxon, extant genus, comprising s ...
. Since the extinction of both wild horses and
aurochs The aurochs (''Bos primigenius''; or ; pl.: aurochs or aurochsen) is an extinct species of Bovini, bovine, considered to be the wild ancestor of modern domestic cattle. With a shoulder height of up to in bulls and in cows, it was one of t ...
en, and the near-extinction of the European bison, domesticated livestock seems to have assumed this role and replaced their extinct relatives. With the abolition of traditional pasturage in town
commons The commons is the cultural and natural resources accessible to all members of a society, including natural materials such as air, water, and a habitable Earth. These resources are held in common even when owned privately or publicly. Commons ...
beginning in the 16th century, however, and the rearing of livestock in factory farms as a consequence of agricultural intensification, these large herbivores are now largely absent from the landscape. Additionally, while research indicates the widespread existence of half-open
savanna A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) biome and ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach th ...
ecosystems during Europe's prehistory, shaped and maintained by
megafauna In zoology, megafauna (from Ancient Greek, Greek μέγας ''megas'' "large" and Neo-Latin ''fauna'' "animal life") are large animals. The precise definition of the term varies widely, though a common threshold is approximately , this lower en ...
, this is no more the case. Nowadays, the landscape in many parts of Europe is marked by closed-canopy forest, often intensively managed, coupled with agricultural fields and urban spaces, with little else and few transitional zones such as mantle and fringe vegetation. As a result, European wild apple today lacks both suitable habitats and dispersal opportunities, resulting in a lack of successful regeneration.


Gallery

File:Malus sylvestris (villeple) epler.jpg, Ripe crab apples on the branch File:Malus sylvestris sl8.jpg, Bark File:Malus sylvestris sl3.jpg, Flowers File:Malus sylvestris - Botanical Garden in Kaisaniemi, Helsinki - DSC03467.JPG, Mature tree File:Disintegrating crab apple tree at Furzy Brow, New Forest - geograph.org.uk - 141960.jpg, Ancient tree File:Abgefallene Holzäpfel.jpg, Ripe fruit on the ground


See also

* Wood-pasture hypothesis * Pleistocene megafauna * '' Pyrus pyraster'' * '' Malus crescimannoi'' * '' Malus orientalis'' * '' Malus trilobata'' * '' Malus florentina''


References


Further reading

* M.H.A. Hoffman, List of names of woody plants, Applied Plant Research, Boskoop 2005. * RHS dictionary of gardening, 1992


External links


USDA Plants Profile for ''Malus sylvestris''

Jepson Manual (JM93) treatment of ''Malus sylvestris''
— ''introduced species in California''. *
EUFORGEN species page: ''Malus sylvestris''
Information, distribution and related resources. {{Authority control sylvestris Crabapples Flora of Europe Flora of Spain Flora of Ukraine Taxa named by Philip Miller