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A sand forest is a type of rare
subtropical The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Geographical z ...
forest A forest is an area of land dominated by trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, and ecological function. The United Nations' ...
region, distinctive due to its unique combination of plant (often rare plant) and animal species, and their restriction to ancient coastal dunes. Sand forests are found in
Maputaland Maputaland is a natural region of Southern Africa. It is located in the northern part of the province of KwaZulu-Natal, South Africa between Eswatini and the coast. In a wider sense it may also include the southernmost region of Mozambique. The bi ...
in South Africa, as well as parts of the
Amazon basin The Amazon basin is the part of South America drained by the Amazon River and its tributaries. The Amazon drainage basin covers an area of about , or about 35.5 percent of the South American continent. It is located in the countries of Bolivi ...
in
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
,
Peru , image_flag = Flag of Peru.svg , image_coat = Escudo nacional del Perú.svg , other_symbol = Great Seal of the State , other_symbol_type = Seal (emblem), National seal , national_motto = "Fi ...
, and
Colombia Colombia (, ; ), officially the Republic of Colombia, is a country in South America with insular regions in North America—near Nicaragua's Caribbean coast—as well as in the Pacific Ocean. The Colombian mainland is bordered by the Car ...
. They are typically composed of clay soils and nutrient poor white sands. These forests are commonly referred to as "sand forests" in the South African regions. However, in Mozambique, they are known as the "Lucauati forests". In the Amazon, they are colloquially known as ''
campinarana Campinarana (NT0158, ), also called Rio Negro Campinarana, is a neotropical ecoregion in the Amazon biome of the north west of Brazil and the east of Colombia that contains vegetation adapted to extremely poor soil. It includes savanna, scrub an ...
s'' and ''
campinas Campinas (, ''Plains'' or ''Meadows'') is a Brazilian municipality in São Paulo State, part of the country's Southeast Region. According to the 2020 estimate, the city's population is 1,213,792, making it the fourteenth most populous Brazilian ...
''. Few studies have been carried out on the sand forests. Those studies that have been conducted, the majority of the research has been on the plant diversity within these forests.


Origins

There have been no comprehensive studies undertaken in the white sand forests. A full understanding of the process in which they have emerged is lacking. However, it is thought that the sand forests are the fragments of coastal dunes which were separated from the ocean over millions of years as the shoreline and water level slowly shifted. Researchers have speculated that these forest systems have been related to the
Late Pleistocene The Late Pleistocene is an unofficial Age (geology), age in the international geologic timescale in chronostratigraphy, also known as Upper Pleistocene from a Stratigraphy, stratigraphic perspective. It is intended to be the fourth division of ...
and
Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togethe ...
dry periods. Podsolization is a process in which the upper layer of soil becomes acidic due to the leaching of nutrients. This process combined with the lack of nutrients in the upper horizon results in only white sand being left behind. Because sandy soils can result from a number of different processes ( nutrient leaching, tectonic activity, river dynamics, etc.), the sand forests are unique and, oftentimes, different from sand forests found elsewhere.


Characteristics


Biodiversity

Sand forests are well known for their unique biodiversity and high levels of
endemism 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 date there have been more than 2,500 species of vascular plants and of those, 230 species are endemic within the Maputaland region. Distinct
sclerophyll Sclerophyll is a type of vegetation that is adapted to long periods of dryness and heat. The plants feature hard leaf, leaves, short Internode (botany), internodes (the distance between leaves along the stem) and leaf orientation which is paral ...
y, or vegetation with thickened, hardened foliage that slows moisture loss, is a characteristic of a vast number of the plants found in this region. Due to nutrient and water restrictions, the vegetation that grows in the sand forests is very specialized. Many of the trees and shrubs have evolved anti-herbivorous defenses. Herbivores seemed to favor vegetation of plants in clay soils over the white sandy soils. In no presence of herbivores however, the clay soil vegetation survived just as well as white sand specialists, but grew much taller and produced more leaf area.


Decomposition

Sand forests have a thick
humus In classical soil science, humus is the dark organic matter in soil that is formed by the decomposition of plant and animal matter. It is a kind of soil organic matter. It is rich in nutrients and retains moisture in the soil. Humus is the Lati ...
layer due to the extremely low decomposition rates. Some sandy forests have even been recorded to have more than a meter of leaf litter on top of the white sand. Multiple reasons exist as to why there is such a large accumulation of organic matter including high acidity of the soil, high content of toxic compounds in the litter, and low nutrient quality of the litter.


Vegetation

It is very common to find
epiphytes 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 ...
, such as the wiry orchid (''
Microcoelia ''Microcoelia'' is a genus of orchids native to sub-Saharan Africa as well as to Madagascar and other islands of the Indian Ocean The Indian Ocean is the third-largest of the world's five oceanic divisions, covering or ~19.8% of the water ...
exilis''), and other lichens growing on the trees. They derive their moisture and nutrients from the air, rain, and debris that has accumulated over time. In contrast to
tropical rainforests Tropical rainforests are rainforests that occur in areas of tropical rainforest climate in which there is no dry season – all months have an average precipitation of at least 60 mm – and may also be referred to as ''lowland equator ...
, campinas are "reduced in biomass and have relatively high light penetration." Due to nutrient deficiencies in campina soils, "shrubs and small trees typically have a dwarfed and rachitic aspect with reduced quantities of foliage" with many of the species being
perennials 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 widel ...
and
evergreen In botany, an evergreen is a plant which has foliage that remains green and functional through more than one growing season. This also pertains to plants that retain their foliage only in warm climates, and contrasts with deciduous plants, which ...
s


List of flora

*
Clusiaceae The Clusiaceae or Guttiferae Juss. (1789) (''nom. alt. et cons.'' = alternative and valid name) are a family of plants including 13 genera and ca 750 species. Several former members of Clusiacae are now placed in Calophyllaceae and Hypericaceae. ...
*''
Pachira ''Pachira'' is a genus of tropical trees distributed in Central and South America, Africa and India. They are classified in the subfamily Bombacoideae of the family Malvaceae. Previously the genus was assigned to Bombacaceae. Prior to that the g ...
brevipes'' *Sali (''
Tetragastris ''Tetragastris'' is a genus of plants in family Burseraceae The Burseraceae are a moderate-sized family of 17-19 genera and about 540 species of flowering plants. The actual numbers differ according to the time period in which a given source i ...
panamensis'') *'' Eperua'' spp. *''
Ocotea ''Ocotea'' is a genus of flowering plants belonging to the family Lauraceae. Many are evergreen trees with lauroid leaves. There are over 520 species currently accepted within the genus, distributed mostly in tropical and subtropical areas of t ...
'' spp. *''
Hortia ''Hortia'' is a genus of plants in family Rutaceae The Rutaceae is a family, commonly known as the rue
'' spp. *''
Inga ''Inga'' is a genus of small tropical, tough-leaved, nitrogen-fixing treesElkan, Daniel. "Slash-and-burn farming has become a major threat to the world's rainforest" ''The Guardian'' 21 April 2004 and shrubs, subfamily Mimosoideae. ''Inga''s l ...
'' spp. *'' Ferdinandusa chlorantha'' *''
Euterpe catinga ''Euterpe catinga'' is a palm species in the genus '' Euterpe''. It is found in forests of a dry, sandy soil and very peculiar vegetation, known as catinga forests or Campinarana in northern South America ( Guyana, Venezuela, Colombia, Ecuador, ...
'' *''
Dendropanax ''Dendropanax'' is a genus of flowering plants in the family Araliaceae, consisting of 92 species of evergreen trees and shrubs, first described by Joseph Decaisne & Jules Émile Planchon in 1854.Decaisne, J. & Planchon, J.E. (1854) Rev. Hort. ar ...
umbellatus'' *''Stigonema tomentosum'' *False tamboti (''Cleisanthus schlechteri'') *False white ash (''Pseudobersama mossambicensis'') * Flat-crown (''Albizia adianthifolia'') *Galla-plum (''
Haplocoelum ''Haplocoelum'' is a genus of plants in the family Sapindaceae The Sapindaceae are a family of flowering plants in the order Sapindales known as the soapberry family. It contains 138 genera and 1858 accepted species. Examples include horse che ...
gallense'') *Glossy flat-bean (''
Dalbergia ''Dalbergia'' is a large genus of small to medium-size trees, shrubs and lianas in the pea family, Fabaceae, subfamily Faboideae. It was recently assigned to the informal monophyletic ''Dalbergia'' clade (or tribe): the Dalbergieae. T ...
nitidula'') *Green thorn (''
Balanites maughamii ''Balanites maughamii'' (manduro, torchwood, af, Groendoring, zu, Ugobandlovu) is a protected tree in South Africa. Groendoring, a community outside Asab Asab ( Nama: ''new thing'') is a settlement in the ǁKaras Region of southern Namibia. ...
'') *Green-apple (''
Monodora ''Monodora'' is a genus of plant in family Annonaceae The Annonaceae are a family of flowering plants consisting of trees, shrubs, or rarely lianas commonly known as the custard apple family or soursop family. With 108 accepted genera and abou ...
junodii'') *Large cluster-pear (''
Uvaria ''Uvaria'' is a genus of flowering plants in the family Annonaceae. The generic name ''uvaria'' is derived from the Latin ''uva'' meaning grape, likely because the edible fruit of some species in the genus resemble grapes. Circumscription Specie ...
lucida'' subsp. ''virens'') *Lavender fever-berry (''
Croton gratissimus ''Croton gratissimus'' (commonly known as lavender croton or lavender fever berry), is a tropical African shrub or small tree with corky bark, growing to 8 m and belonging to the family of Euphorbiaceae or spurges. Young twigs are slender an ...
'') *Lavender-leaved croton ('' Croton pseudopulchellus'') *Lebombo wattle (''
Newtonia hildebrandtii ''Newtonia hildebrandtii'', the Lebombo wattle ( af, Lebombowattel, zu, Umfomothi), is a medium-sized tree native to eastern Africa. It is a protected tree in South Africa. Description ''Newtonia hildebrandtii'' is a medium-sized tree growing ...
'') *Lowveld milkberry (''
Manilkara ''Manilkara'' is a genus of trees in the family Sapotaceae. They are widespread in tropical and semitropical locations, in Africa, Madagascar, Asia, Australia, and Latin America, as well as various islands in the Pacific and in the Caribbean. A ...
mochisia'') *Marsh fever-berry (''Croton steenkampianus'') *Mozambique coffee (''
Coffea ''Coffea'' is a genus of flowering plants in the Family (biology), family Rubiaceae. ''Coffea'' species are shrubs or small trees native to tropical and southern Africa and tropical Asia. The seeds of some species, called coffee beans, are use ...
racemosa'') *Pink lannea (''
Lannea ''Lannea'' is a genus of plants in the family Anacardiaceae The Anacardiaceae, commonly known as the cashew family or sumac family, are a family of flowering plants, including about 83 genera with about 860 known species. Members of the Anaca ...
antiscorbutica'') *Pod mahogany (
Afzelia quanzensis ''Afzelia quanzensis'' (pod mahogany, af, peulmahonie, ve, mutokota, zu, inkehli) is a protected tree in South Africa. This deciduous tree can grow up to 20m in height, and has smooth, grey bark, which can flake in irregular patches. It has glo ...
) *Quar (''
Psydrax ''Psydrax'' is a genus of flowering plants in the family Rubiaceae. It consists of trees, shrubs, and a few lianas in the paleotropics. Taxonomy The genus was named by Joseph Gaertner in 1788 in his book, ''De Fructibus et Seminibus Plantarum'' ...
obovata'') *Quiver-leaf fig (''
Ficus ''Ficus'' ( or ) is a genus of about 850 species of woody trees, shrubs, vines, epiphytes and hemiepiphytes in the family Moraceae. Collectively known as fig trees or figs, they are native throughout the tropics with a few species extending in ...
tremula'') *Red-hair bush (''
Lasiodiscus ''Lasiodiscus'', commonly known as red-hair bushes, is a small plant genus in the family Rhamnaceae. It is endemic to Africa and its adjacent islands. Description The small trees have opposite, often asymmetric leaves. As with ''Colubrina'', th ...
pervillei'' subsp. ''pervillei'') *Red-heart tree (''Hymenocarida ulmoides'') *Saddle-pod ('' Wrightia natalensis'') *Sand cabbage tree (''
Cussonia ''Cussonia'' is a genus of plants of the family Araliaceae, which is native to the Afrotropics. It originated in Africa and has its center of distribution in South Africa and the Mascarene Islands. Due to their striking habit, they are a conspi ...
arenicola'') *Sand canary-berry ('' Suregada zanzibariensis'') *Sand forest gardenia ('' Hyperacanthus microphyllus'') *Sand ivory (''
Berchemia ''Berchemia'' is a genus of plants in the family Rhamnaceae, named after Dutch botanist Berthout van Berchem. They are climbing plants or small to medium-sized trees that occur in Africa, Asia and the Americas The Americas, which are s ...
'' sp. nov.) *Sand forest hum-num (''
Carissa tetramera ''Carissa tetramera'', the sand num-num, is a plant in the dogbane family Apocynaceae. The specific epithet ''tetramera'' means "four parts", referring to the flower. Description ''Carissa tetramera'' grows as a shrub up to tall. Its strong-sc ...
'') *Sand knobwood (''Zanthoxylon leprieuri'') *Sherbet tree (''
Dialium ''Dialium'' is a genus of flowering plants in the family Fabaceae, subfamily Dialioideae. Velvet tamarind is a common name for several species. Species , The Plant List accepts the following species: *'' Dialium angolense'' Oliv. *''Dialium au ...
schlechteri'') *Sneezewood (''
Ptaeroxylon obliquum ''Ptaeroxylon obliquum'' is the botanical name for the sneezewood tree. It is native to Southern Africa, including South Africa, Zimbabwe, and Mozambique. It is the only species in the genus ''Ptaeroxylon''. Background ''Ptaeroxylon obliquum' ...
'') *Stink bushwillow (''
Pteleopsis myrtifolia ''Pteleopsis myrtifolia'' ('Ptelea' = elm, 'opsis' = resembling, 'myrtifolia' = leaves like those of the myrtle) is one of some 10 African species in this genus in the family of Combretaceae. It is the only ''Pteleopsis'' species to occur in Sout ...
'') *Swazi ordeal tree (''
Erythrophleum lasianthum ''Erythrophleum lasianthum'', the Maputaland ordeal tree, is a tree with a localized range in the endangered lowland forests of southeastern Africa. Range It is found from False Bay to Maputaland in northern KwaZulu-Natal province of South Afri ...
'') *Tonga-kerrie ( Cladostemon kirkii) *Water ironplum (''
Drypetes ''Drypetes'' is a plant genus of the family Putranjivaceae, in the order Malpighiales. It was previously in the family Euphorbiaceae, tribe Drypeteae, and was the sole pantropical zoochorous genus of the family. The genus comprises about 200 ...
arguta'') *Wild mandarin (''Toddaliopsis bremekampii'') *Wild mango ('' Cordyla africana'') *Zulu coshwood (''
Cola Cola is a carbonated soft drink flavored with vanilla, cinnamon, citrus oils and other flavorings. Cola became popular worldwide after the American pharmacist John Stith Pemberton invented Coca-Cola, a trademarked brand, in 1886, which was imita ...
greenwayi'') *Zulu loquat ('' Oxyanthus latifolius'')


Hydrology

Sand forests have vegetation
gradient In vector calculus, the gradient of a scalar-valued differentiable function of several variables is the vector field (or vector-valued function) \nabla f whose value at a point p is the "direction and rate of fastest increase". If the gradi ...
s that closely coincide with saturation gradients. These saturation gradients are affected by rainfall and
topography Topography is the study of the forms and features of land surfaces. The topography of an area may refer to the land forms and features themselves, or a description or depiction in maps. Topography is a field of geoscience and planetary sci ...
and, therefore, the vegetation is also affected by these same factors. Vegetation lower to the ground tends to congregate in areas that have been saturated for the longest period of time.
Blackwater river A blackwater river is a type of river with a slow-moving channel flowing through forested swamps or wetlands. As vegetation decays, tannins leach into the water, making a transparent, acidic water that is darkly stained, resembling black te ...
s (e.g. the Rio Negro) commonly begin in these sand forests due to their accumulation of humic matter that is easily washed downstream, particularly after heavy rainfall.


Birds

The majority of bird species found in sand forests is known to prefer sand forest habitat and are found rarely, if at all, in other types of habitats. There have been no studies, however, evaluating the habitat specialization of sand forest birds to determine whether these habitats have unique bird fauna. The majority of these birds are small ground dwellers. They often forage in the underbrush to find food because of the thick organic layers found in the sand forests. Their sharp beaks allow them to break through the thick coating of fruits and obtain the seeds inside.


List of bird species

* '' Attila citriniventris'' * '' Claravis pretiosa'' * '' Cnemotriccus duidae'' * '' Crypturellus strigilosus'' * '' Conopias parva'' * '' Crytpturellus duidae'' * '' Deconychura longicauda'' * '' Dixiphia pipra'' * '' Galbula dea'' * '' Hemitriccus minimus'' * ''
Herpsilochmus gentryi The ancient antwren (''Herpsilochmus gentryi'') is a species of tropical bird in the family Thamnophilidae. It is primarily found in ''terra firme'' forests of northern Peru and southeastern Ecuador. This species was described in 1998 and named a ...
'' * '' Heterocercus aurantiivertex'' * '' Hypocnemis hypoxantha'' * '' Lepidocolaptes albolineatus'' * '' Megastictus margaritatus'' * '' Myrmeciza castanea'' * '' Myrmotherula leucophthalma'' * '' Neopelma chrysocephalum'' * '' Neopipo cinnamomea'' * '' Notharchus ordii'' * '' Percnostola arenarum'' * '' Pithys castanea'' * '' Platyrinchus saturatus'' * ''
Polioptila ''Polioptila'' is a genus of small insectivorous birds in the family Polioptilidae. They are found in North and South America. The genus ''Polioptila'' was introduced by the English zoologist Philip Sclater in 1855. Although he listed several m ...
'' sp. nov. * '' Polytmus theresiae'' * '' Ramphotrigon ruficauda'' * ''
Sclerurus rufigularis The short-billed leaftosser (''Sclerurus rufigularis'') is a species of bird in the family Furnariidae. It is found in Bolivia, Brazil, Colombia, Ecuador, French Guiana, Guyana, Peru, Suriname, and Venezuela. Its natural habitat is subtropical o ...
'' * '' Tachyphonus phoenicius'' * '' Trogon rufus'' * ''
Xenopipo atronitens The black manakin (''Xenopipo atronitens'') is a species of bird in the family Pipridae. It is found in Bolivia, Brazil, Colombia, French Guiana, Guyana, Peru, Suriname, and Venezuela. Its natural habitats are subtropical or tropical moist lowla ...
'' * '' Xipholena punicea'' * '' Zimmerius villarejoi''


The role of fire

The exact role of fires in sand forests is not known and most studies disagree on the matter. However, evidence has shown that fires can help increase species diversity, particularly those that are specialized for such areas. For example, fire tolerant species may be more likely to colonize here. Historically speaking, some sand forests may have been the result of anthropogenic burning done by indigenous people. They would burn
swath A small waterplane area twin hull, better known by the acronym SWATH, is a catamaran design that minimizes hull cross section area at the sea's surface. Minimizing the ship's volume near the surface area of the sea, where wave energy is locate ...
s of sand forest to obtain better hunting and farming grounds. Fires of any kind can have both positive and negative effects on the ecosystem. In some instances, a fire can cause one or just a few species to predominate when the forest begins to regenerate. However, fire can also promote more diversity within the regenerated forest.


Threats

Because sand forests are isolated in small patches, they have been mostly protected from the
destruction of the Amazon rainforest The Amazon rainforest is the largest rainforest in the world, covering an area of 3,000,000 km2 (2,316,612.95 square miles). It represents over half of the planet's rainforests and comprises the largest and most biodiverse tract of tropi ...
. However, they are still at risk of disturbance. Sand forests are extremely sensitive to destruction due to the harsh growth and survival conditions. Since the conditions are so tough to thrive in once an ecosystem declines, it may take hundreds of years for it to develop again. Some sand forests overlap with protected areas and indigenous lands, but most do not. Although they are not immediately threatened, sand forests are still in danger from ever-increasing
deforestation Deforestation or forest clearance is the removal of a forest or stand of trees from land that is then converted to non-forest use. Deforestation can involve conversion of forest land to farms, ranches, or urban use. The most concentrated d ...
, hydroelectric projects, and mineral extractions.


Conservation

Peru has actively incorporated sand forests in three protected areas. Authors who have done research in campinas have also made recommendations for conservation in their work.


See also

*
Southern African Sand Forest Southern African Sand Forest is a sand forest, or a subtropical forest plant community of the tropical and subtropical dry broadleaf forests biome. It grows on ancient sand dunes in northern KwaZulu-Natal and southern Mozambique. In South Africa ...


References

{{reflist, 2 Tropical and subtropical dry broadleaf forests Amazon basin