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Sclerophyll is a type of vegetation that is adapted to long periods of dryness and heat. The plants feature hard
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 ...
, short internodes (the distance between leaves along the stem) and leaf orientation which is parallel or oblique to direct sunlight. The word comes from the Greek ''sklēros'' (hard) and ''phyllon'' (leaf). The term was coined by A.F.W. Schimper in 1898 (translated in 1903), originally as a synonym of
xeromorph A xerophyte (from Ancient Greek language, Greek ξηρός ''xeros'' 'dry' + φυτόν ''phuton'' 'plant') is a species of plant that has adaptations to survive in an environment with little liquid water, such as a desert such as the Sahara or pl ...
, but the two words were later differentiated. Sclerophyllous plants occur in many parts of the world, but are most typical of areas with low rainfall or seasonal droughts, such as Australia, Africa, and western North and South America. They are prominent throughout
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
, parts of Argentina, the Cerrado biogeographic region of
Bolivia , image_flag = Bandera de Bolivia (Estado).svg , flag_alt = Horizontal tricolor (red, yellow, and green from top to bottom) with the coat of arms of Bolivia in the center , flag_alt2 = 7 × 7 square p ...
, Paraguay and Brazil, and in the Mediterranean biomes that cover the
Mediterranean Basin In biogeography, the Mediterranean Basin (; also known as the Mediterranean Region or sometimes Mediterranea) is the region of lands around the Mediterranean Sea that have mostly a Mediterranean climate, with mild to cool, rainy winters and w ...
, California, Chile, and the Cape Province of South Africa. In the
Mediterranean basin In biogeography, the Mediterranean Basin (; also known as the Mediterranean Region or sometimes Mediterranea) is the region of lands around the Mediterranean Sea that have mostly a Mediterranean climate, with mild to cool, rainy winters and w ...
,
holm oak Holm oak may refer to: * '' Quercus ilex'', tree native to South and Southeast Europe and parts of France * '' Quercus rotundifolia'', tree native to the Iberian Peninsula and Northwest Africa * ''Quercus agrifolia ''Quercus agrifolia'', the Cal ...
, cork oak and
olive The olive, botanical name ''Olea europaea'', meaning 'European olive' in Latin, is a species of small tree or shrub in the family Oleaceae, found traditionally in the Mediterranean Basin. When in shrub form, it is known as ''Olea europaea'' ...
s are typical hardwood trees. In addition, there are several species of pine under the trees in the vegetation zone. The shrub layer contains numerous herbs such as rosemary, thyme and lavender. In relation to the potential natural vegetation, around 2% of the earth's land surface is covered by sclerophyll woodlands, and a total of 10% of all plant species on earth live there.


Description

Sclerophyll woody plants are characterized by their relatively small, stiff, leathery and long-lasting leaves. The sclerophyll vegetation is the result of an adaptation of the flora to the summer dry period of a Mediterranean-type climate. Plant species with this type of adaptation tend to be evergreen with great longevity, slow growth and with no loss of leaves during the unfavorable season. To note, "sclerophyll" and "evergreen" are not synonymous. As a result, the thickets that make up these ecosystems are of the persistent evergreen type, in addition to the predominance of plants, even
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 ...
ones, with "hard" leaves, which are covered by a thick leathery layer called the
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
, that prevents water loss during the dry season. The aerial and underground structures of these plants are modified to make up for water shortages that may affect their survival. The name sclerophyll derives from the highly developed sclerenchyma from the plant, which is responsible for the hardness or stiffness of the leaves. This structure of the leaves inhibits transpiration and thus prevents major water losses during the dry season. Most of the plant species in the sclerophyll zone are not only insensitive to summer drought, they have also used various strategies to adapt to frequent wildfires, heavy rainfall and
nutrient A nutrient is a substance used by an organism to survive, grow, and reproduce. The requirement for dietary nutrient intake applies to animals, plants, fungi, and protists. Nutrients can be incorporated into cells for metabolic purposes or excret ...
deficiencies.


Ecology

The
type Type may refer to: Science and technology Computing * Typing, producing text via a keyboard, typewriter, etc. * Data type, collection of values used for computations. * File type * TYPE (DOS command), a command to display contents of a file. * Ty ...
of sclerophyllic trees in the
Palearctic The Palearctic or Palaearctic is the largest of the eight biogeographic realms of the Earth. It stretches across all of Eurasia north of the foothills of the Himalayas, and North Africa. The realm consists of several bioregions: the Euro-Sibe ...
flora region include the holm oak (''
Quercus ilex ''Quercus ilex'', the evergreen oak, holly oak or holm oak is a large evergreen oak native to the Mediterranean region. It is a member of the ''Ilex'' section of the genus, with acorns that mature in a single summer. Description An evergreen tr ...
''), myrtle (''
Myrtus communis ''Myrtus communis'', the common myrtle or true myrtle, is a species of flowering plant in the myrtle family Myrtaceae. It is an evergreen shrub native to southern Europe, North Africa, Western Asia, Macaronesia, and the Indian Subcontinent, and a ...
''), strawberry tree ('' Arbutus unedo''), wild olive (''
Olea europaea The olive, botanical name ''Olea europaea'', meaning 'European olive' in Latin, is a species of small tree or shrub in the family Oleaceae, found traditionally in the Mediterranean Basin. When in shrub form, it is known as ''Olea europaea'' ...
''), laurel ('' Laurus nobilis''), mock privet (''
Phillyrea latifolia ''Phillyrea latifolia'', commonly known as green olive tree or mock privet, is a species of tree in the family Oleaceae. It is native to the Mediterranean Basin In biogeography, the Mediterranean Basin (; also known as the Mediterranean R ...
''), the Italian buckthorn (''
Rhamnus alaternus ''Rhamnus alaternus'' is a species of flowering plant in the buckthorn family Rhamnaceae, known by the common names Italian buckthorn or Mediterranean buckthorn. It is a hardy medium-sized evergreen shrub with fragrant flowers. Etymology The sp ...
''), etc. * In central and southern California, the coastal hills are covered in sclerophyll vegetation known as chaparral. The flora of this ecoregion also includes tree species Scrub oak (''Quercus dumosa''), California buckeye (''Aesculus californica''), San Gabriel Mountain liveforever (''Dudlea densiflora''), Catalina mahogany (''Cercocarpus traskiae''), and the threatened jewelflower (''Streptanthus albidus'' ssp. ''Peramoenus''). (material included verbatim under th
CC BY-SA 3.0 license
/ref> * In South Africa, in the Cape region, there are Mediterranean open forests known as
fynbos Fynbos (; meaning fine plants) is a small belt of natural shrubland or heathland vegetation located in the Western Cape and Eastern Cape provinces of South Africa. This area is predominantly coastal and mountainous, with a Mediterranean clim ...
. The abundance of endemics is so extraordinary (68% of the 8600 vascular plant species in the area) that the South African sclerophyll area, the cape flora, forms the smallest of the six flora kingdoms on earth. Plants include ''
Elegia Elegia may refer to: * The Latin term for "elegy" * ''Elegia'' (moth), a snout moth genus in subfamily Phycitinae * ''Elegia'' (plant), a South African plant genus in family Restionaceae * , 1979 Polish film directed by Paweł Komorowski * Elegia ...
'', ''
Thamnochortus ''Thamnochortus'' is a group of plants in the Restionaceae described as a genus in 1767.Bergius, Peter Jonas. 1767. Descriptiones Plantarum ex Capite Bonae Spei 353 The entire genus is endemic to Cape Province in South Africa. ; Species ; forme ...
'', and '' Willdenowia'' and proteas such as king protea ('' Protea cynaroides'') and blushing bride ('' Serruria florida''). * In most of Australia, sclerophyll vegetation such as eucalyptus trees,
melaleuca ''Melaleuca'' () is a genus of nearly 300 species of plants in the myrtle family, Myrtaceae, commonly known as paperbarks, honey-myrtles or tea-trees (although the last name is also applied to species of '' Leptospermum''). They range in size ...
s, banksias,
callistemon ''Callistemon'' is a genus of shrubs in the family Myrtaceae, first described as a genus in 1814. The entire genus is endemic to Australia but widely cultivated in many other regions and naturalised in scattered locations. Their status as a se ...
s and grevilleas dominate the mallee and woodland areas of its cities, including those lacking a Mediterranean climate, such as
Sydney Sydney ( ) is the capital city of the state of New South Wales, and the most populous city in both Australia and Oceania. Located on Australia's east coast, the metropolis surrounds Sydney Harbour and extends about towards the Blue Mountain ...
, Melbourne,
Hobart Hobart ( ; Nuennonne/Palawa kani: ''nipaluna'') is the capital and most populous city of the Australian island state of Tasmania. Home to almost half of all Tasmanians, it is the least-populated Australian state capital city, and second-small ...
and Brisbane. * In Chile, south of the desert areas, there is evergreen bushland called
matorral 300px, Springtime in Chilean matorral a few kilometers north of Santiago along the Pan-American Highway Matorral is a Spanish language, Spanish word, along with ''tomillares'', for shrubland, thicket or bushes. It is used in naming and describin ...
. Typical species include Litre ''(Lithraea venenosa)'', Quillay or Soapbark Tree ''( Quillaja saponaria)'', and bromeliads of genus '' Puya''.


Climate

The sclerophyll regions are located in the outer subtropics bordering the temperate zone (also known as the warm-temperate zone). Accordingly, the annual average temperatures are relatively high at 12–24 °C; An average of over 18 °C is reached for at least four months, eight to twelve months it is over 10 °C and no month is below 5 °C on average.
Frost Frost is a thin layer of ice on a solid surface, which forms from water vapor in an above-freezing atmosphere coming in contact with a solid surface whose temperature is below freezing, and resulting in a phase change from water vapor (a gas) ...
and snow occur only occasionally and the
growing season A season is a division of the year marked by changes in weather, ecology, and the amount of daylight. The growing season is that portion of the year in which local conditions (i.e. rainfall, temperature, daylight) permit normal plant growth. Whil ...
lasts longer than 150 days and is in the winter half-year. The lower limit of the moderate annual precipitation is 300mm ( semi-arid climate) and the upper limit 900–1000 mm. Generally, the summers are dry and hot with a dry season of a maximum of seven months, but at least two to three months. The winters are rainy and cool. However, not all regions with sclerophyll vegetation feature the classic Mediterranean climate; parts of eastern Italy, eastern Australia and eastern South Africa, which feature sclerophyll woodlands, tend to have uniform rainfall or even a more summer-dominant rainfall, whereby falling under the
humid subtropical climate A humid subtropical climate is a zone of climate characterized by hot and humid summers, and cool to mild winters. These climates normally lie on the southeast side of all continents (except Antarctica), generally between latitudes 25° and 40° ...
zone (''Cfa''/''Cwa''). Furthermore, other areas with sclerophyll flora would grade to the
oceanic climate An oceanic climate, also known as a marine climate, is the humid temperate climate sub-type in Köppen classification ''Cfb'', typical of west coasts in higher middle latitudes of continents, generally featuring cool summers and mild winters ( ...
(''Cfb''); particularly the eastern parts of the
Eastern Cape The Eastern Cape is one of the provinces of South Africa. Its capital is Bhisho, but its two largest cities are East London and Gqeberha. The second largest province in the country (at 168,966 km2) after Northern Cape, it was formed in ...
province in South Africa, and Tasmania, Victoria and southern New South Wales in Australia.


Distribution

The zone of the sclerophyll vegetation lies in the border area between the
subtropics The subtropical zones or subtropics are geographical and climate zones to the north and south of the tropics. Geographically part of the temperate zones of both hemispheres, they cover the middle latitudes from to approximately 35° north and ...
and the temperate zone, approximately between the 30th and 40th degree of latitude (in the northern hemisphere also up to the 45th degree of latitude). Their presence is limited to the coastal western sides of the continents, but nonetheless can typical in any regions of a continent with scarce annual precipitation or frequent seasonal droughts and poor soils that are heavily leached. The sclerophyll zone often merges into temperate deciduous forests towards the poles, on the coasts also into temperate rainforests and towards the equator in hot
semi-desert A semi-arid climate, semi-desert climate, or steppe climate is a dry climate sub-type. It is located on regions that receive precipitation below potential evapotranspiration, but not as low as a desert climate. There are different kinds of semi-ar ...
s or deserts. The Mediterranean areas, which have a very high biodiversity, are under great pressure from the population. This is especially true for the Mediterranean region since ancient times. Through
overexploitation Overexploitation, also called overharvesting, refers to harvesting a renewable resource to the point of diminishing returns. Continued overexploitation can lead to the destruction of the resource, as it will be unable to replenish. The term app ...
(logging, grazing, agricultural use) and frequent fires caused by people, the original forest vegetation is converted. In extreme cases, the hard-leaf vegetation disappears completely and is replaced by open rock heaths. Some sclerophyll areas are closer to the equator than the Mediterranean zone—for example, the interior of Madagascar, the dry half of
New Caledonia ) , anthem = "" , image_map = New Caledonia on the globe (small islands magnified) (Polynesia centered).svg , map_alt = Location of New Caledonia , map_caption = Location of New Caledonia , mapsize = 290px , subdivision_type = Sovereign st ...
, the lower edge areas of the Madrean pine-oak woodlands of the Mexican highlands between 800 and 1800/2000 m or around 2000 m high plateaus of the Asir Mountains on the western edge of the
Arabian Peninsula The Arabian Peninsula, (; ar, شِبْهُ الْجَزِيرَةِ الْعَرَبِيَّة, , "Arabian Peninsula" or , , "Island of the Arabs") or Arabia, is a peninsula of Western Asia, situated northeast of Africa on the Arabian Plate ...
.


Land use

While the winter rain areas of America, South Africa and
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
, with an unusually large variety of food crops, were ideal gathering areas for hunter gatherers until European colonization, agriculture and
cattle breeding Animal husbandry is the branch of agriculture concerned with animals that are raised for meat, fibre, milk, or other products. It includes day-to-day care, selective breeding, and the raising of livestock. Husbandry has a long history, startin ...
spread in the Mediterranean area since the Neolithic, which permanently changed the face of the landscape. In the sclerophyll regions near the coast, permanent crops such as olive and wine cultivation established themselves; However, the landscape forms that characterize the degenerate shrubbery and shrub heaths
Macchie 220px, Low maquis in Corsica 220px, High ''macchia'' in Sardinia ( , , ) or ( , ; often in Italian; hr, makija; ; ) is a shrubland biome in the Mediterranean region, typically consisting of densely growing evergreen shrubs. Maquis is cha ...
and Garigue are predominantly a result of grazing (especially with goats). In the course of the last millennia, the original vegetation in almost all areas of this vegetation zone has been greatly changed by the influence of humans. Where the plants have not been replaced by vineyards and olive grove (nature), groves, the maquis shrubland, maquis was the predominant form of vegetation on the Mediterranean. The maquis has been degraded in many places to the low shrub heather, the garigue. Many plant species that are rich in aromatic oils belong to both vegetation societies. The diversity of the original sclerophyll vegetation in the world is high to extremely high (3000-5000 species per ha).


Australian bush

Most areas of the Australian continent able to support woody plants are occupied by sclerophyll communities as forests, savannas, or Heath (habitat), heathlands. Common plants include the Proteaceae ( grevilleas, banksias and hakeas), Melaleuca, tea-trees, acacias, boronias, and eucalypts. The most common sclerophyll communities in Australia are savannas dominated by grasses with an overstorey of eucalypts and acacias. Acacia (particularly Acacia aneura, mulga) shrublands also cover extensive areas. All the dominant overstorey acacia species and a majority of the understorey acacias have a scleromorphic adaptation in which the leaves have been reduced to phyllodes consisting entirely of the Petiole (botany), petiole. Many plants of the sclerophyllous woodlands and shrublands also produce leaves unpalatable to herbivores by the inclusion of toxic and indigestible compounds which assure survival of these long-lived leaves. This trait is particularly noticeable in the eucalypt and ''Melaleuca'' species which possess oil glands within their leaves that produce a pungent volatile oil that makes them unpalatable to most browsers. These traits make the majority of woody plants in these woodlands largely unpalatable to domestic livestock. It is therefore important from a grazing perspective that these woodlands support a more or less continuous layer of herbaceous ground cover dominated by grasses. Sclerophyll forests cover a much smaller area of the continent, being restricted to relatively high rainfall locations. They have a eucalyptus overstory (10 to 30 metres) with the understory also being hard-leaved. ''Dry sclerophyll'' forests are the most common forest type on the continent, and although it may seem barren dry sclerophyll forest is highly diverse. For example, a study of sclerophyll vegetation in Seal Creek, Victoria (Australia), Victoria, found 138 species. Even less extensive are ''wet sclerophyll'' forests. They have a taller eucalyptus overstory than dry sclerophyll forests, 30 metres or more (typically Eucalyptus regnans, mountain ash, Eucalyptus delegatensis, alpine ash, Eucalyptus grandis, rose gum, Eucalyptus diversicolor, karri, Eucalyptus obliqua, messmate stringybark, or Eucalyptus viminalis, manna gum, and a soft-leaved, fairly dense understory (tree ferns are common). They require ample rainfall—at least 1000 mm (40 inches).


Evolution

Sclerophyllous plants are all part of a specific environment and are anything but newcomers. By the time of European settlement, sclerophyll forest accounted for the vast bulk of the forested areas. Most of the wooded parts of present-day Australia have become sclerophyll dominated as a result of the extreme age of the continent combined with Indigenous Australians, Aboriginal fire use. Deep weathering of the crust over many millions of years leached chemicals out of the rock, leaving Australian soils deficient in nutrients, particularly phosphorus. Such nutrient deficient soils support non-sclerophyllous plant communities elsewhere in the world and did so over most of Australia prior to European arrival. However such deficient soils cannot support the nutrient losses associated with frequent fires and are rapidly replaced with sclerophyllous species under traditional Aboriginal burning regimens. With the cessation of traditional burning non-sclerophyllous species have re-colonized sclerophyll habitat in many parts of Australia. The presence of toxic compounds combined with a high carbon : nitrogen ratio make the leaves and branches of scleromorphic species long-lived in the litter, and can lead to a large build-up of litter in woodlands. The toxic compounds of many species, notably Eucalyptus species, are volatile and flammable and the presence of large amounts of flammable litter, coupled with an herbaceous understorey, encourages fire. All the Australian sclerophyllous communities are liable to be burnt with varying frequencies and many of the woody plants of these woodlands have developed adaptations to survive and minimise the effects of fire. Sclerophyllous plants generally resist dry conditions well, making them successful in areas of seasonally variable rainfall. In Australia, however, they evolved in response to the low level of phosphorus in the soil—indeed, many native Australian plants cannot tolerate higher levels of phosphorus and will die if fertilised incorrectly. The leaves are hard due to lignin, which prevents wilting and allows plants to grow, even when there is not enough phosphorus for substantial new cell growth.


Regions

These are the biomes or ecoregions in the world that feature an abundance of, or are known for having, sclerophyll vegetation: *Cumberland Plain Woodland *Sydney Sandstone Ridgetop Woodland *Tasmanian dry sclerophyll forests *Aegean and Western Turkey sclerophyllous and mixed forests *California chaparral and woodlands *California coastal sage and chaparral *Chilean Matorral *Mallee Woodlands and Shrublands *Italian sclerophyllous and semi-deciduous forests *Eastern Mediterranean conifer–sclerophyllous–broadleaf forests *Southwest Iberian Mediterranean sclerophyllous and mixed forests *Tyrrhenian–Adriatic sclerophyllous and mixed forests *Canary Islands dry woodlands and forests *Mediterranean acacia–argania dry woodlands *Mediterranean dry woodlands and steppe *Southeastern Iberian shrubs and woodlands *Cyprus Mediterranean forests *Crete Mediterranean forests *Cape Floristic Region *Southern Anatolian montane conifer and deciduous forests *Albany thickets *Northwest Iberian montane forests


See also

*Mediterranean forests, woodlands, and scrub *Chaparral *Fynbos *Maquis shrubland *Garrigue *Kwongan *Matorral *Barren vegetation


References

{{reflist, 2 Mediterranean forests, woodlands, and scrub California chaparral and woodlands Chilean Matorral Mallee Woodlands and Shrublands Ecology Sclerophyll forests