Atlantic Coastal Forest of Brazil (Southern Bahia)
These forests represent one of the last strongholds of Atlantic lowland forest in eastern Brazil. They are structurally similar to Amazonian forest although compared with Amazonia there are far fewer buttressed roots. They are also taxonomically similar but include a high degree of endemism in their own right. Another interesting feature is the large number of primitive plants and are especially well known for their primitive bambusoid grasses. Stratification appears to vary but at least two tree layers have been recognized in southern Bahia including emergent and canopy layers. The most frequent trees include Diploon cuspidatum, Eriotheca macrophylla and the endemic Macrolobium latifolium (Caesalpiniaceae) and Rinorea bahiensis (Violaceae). Other endemic trees include Actinostemon concolor, Mabea glaziouvii, Pausandra morisiana (Euphorbiaceae), Affonsea densiflora, Calliandra bella, Inga blanchetiana, Stryphnodendron pulcherrimum (Mimosaceae), Andira nitida, Swartzia apetala, Vataireopsis araoba (Fabaceae), Aniba intermedia, Licaria bahiana (Lauraceae), Arapatiella emarginata, Chamaecrista bahiae, Harleyodendron unifoliolatum (Caesalpiniaceae), Attalea funifera, Euterpe edulis, Geonoma pauciflora (Arecaceae), Banara kuhlmannii, Carpotroche brasiliense, Casearia bahiensis (Flacourtiaceae), Couepia belemii, Hirtella angustifolia, Licania bahiensis (Chrysobalanaceae), Dulacia papillosa (Olacaceae), Elvasia tricarpellata (Ochnaceae), Eschweilera alvimii (Lecythidaceae), Guarea blanchetii, Trichilia ramalhoi (Meliaceae), Hornschuchia brytrophe (Annonaceae), Hortia arborea, Pilocarpus riedelianus (Rutaceae), Jacaratia heptaphylla (Caricaceae), Lacistema robustum (Lacistemataceae), Manilkara longifolia, Micropholis gardneriana (Sapotaceae), Miconia octopetala, Mouriri bahiensis, Tibouchina elegans (Melastomataceae), Paypayrola blanchetiana (Violaceae), Sloanea alnifolia (Elaeocarpaceae), Stephanopodium blanchetianum (Dichapetalaceae) and Virola officinalis (Myristicaceae). Tree diversity is extremely high and comparable to some of the most species rich forests in Amazonia. The epiphytic flora is also very rich. A recent study showed that up to 80% of trees supported at least some epiphytes on their trunks. These include many species of fern and Araceae but it is the Bromeliaceae that dominate the epiphytic flora in South Bahia, especially those that store water in leaf rosettes. Conspicuous are the ‘large tank’ species such as Aechmea conifera and Hohenbergia blanchetti. Other smaller but more abundant species include Aechmea lingulata, Araeococcus parvifolius and Guzmannia lingulata. Others epiphytes include the endemic Heteropsis oblongifolia, Philodendron williamsii, Rhodospatha latifolia (Araceae) and Rhipsalis pachyptera (Cactaceae). An unusual epiphyte is Codonanthe uleana (Gesneriaceae) with its large root-ball inhabited by ants. Lianas are less conspicuous. Studies show that only about 37% of trees support them. Nevertheless, there are a number of endemic species such as Forsteronia leptocarpa and Mandevilla permixta (Apocynaceae). The shrub layer includes species such as the endemic Aphelandra blanchetiana, Ruellia affinis (Acanthaceae), Baccharis singularis (Asteraceae) and Erythroxylum columbinum (Erythroxylaceae). The herb layer also includes many endemic species such as Anthurium bellum, Asterostigma riedelianum, Dracontioides descisens, Zomicarpa steigeriana (Araceae), Becquerelia clarkei, Pleurostachys gaudichaudii (Cyperaceae), Begonia subacida (Begoniaceae), Dichorisandra leucophthalmos (Commelinaceae) and Justicia symphyantha (Acanthaceae).
Atlantic Interior Forest of Paraguay
These forests rarely form large continuous stands but tend to alternate with areas of savanna. They are also influenced along their margins by other floras including the Chaco flora in the west, the Cerrado flora in the north and Pampas flora in the south. Nevertheless, they still constitute a major centre of endemism particularly for bird species. They typically comprise three well-differentiated tree strata and the canopy or upper stratum can reach heights of up to 40 m. The main canopy trees include Anadenanthera colubrina, Apuleia leiocarpa, Cordia trichotoma, Patagonula americana, Peltophorum dubium, Pterogyne nitens and the endemic Balfourodendron riedelianum (Rutaceae) and Cedrela fissilis (Meliaceae), Many of these have important commercial value. An intermediate tree stratum ranges in height from 10-20 m and commonly includes Holocalyx balansae. Other under storey trees include Cecropia pachystachya, Chrysophyllum gonocarpum, Diatenopteryx sorbifolia, Guarea guidonia, Hennecartia omphalandra, Inga affinis, Jacaratia corumbensis, Nectandra angustifolia, Pilocarpus pennatifolius, Plinia rivularis, Sorocea bonplandii, Trichilia catigua and the endemic Campomanesia xanthocarpa (Myrtaceae). Many of these under story trees produce edible fruits and provide an important resource for local fauna. Common shrub layer species include Piper hispidum and Piper medium together with bamboos such as Chesquea ramosissima, Merostachys claussenii and Olyra micrantha. At ground level there are various herbs and terrestrial ferns. The latter include Adianthopsis radiata, Blechnum brasiliense, Didymochlaena truncatula and Pteris denticulata. Other endemic species of these forests include Annona amambayensis (Annoniaceae), Aspidosperma polyneuron (Apocynaceae), Astronium fraxinifolium (Anacardiaceae), Cabralea canjerana (Meliaceae), Euterpe edulis (Arecaceae), Myrocarpus frondosus and Tabebuia heptaphylla (Bignoniaceae). Another important feature are the occasional almost exclusive stands of the arborescent ferns species Alsophila cuspidata and Cyathea atrovirens. These can grow to heights of up to 7 m and are often associated with slopes or streams or certain soil conditions. Associated species may include Cecropia pachystachya, Luehea divaricata and Peltophorum dubium. These areas often have an extremely rich epiphytic flora of ferns, cacti and orchids giving them an almost ‘magic’ appearance.
Atlantic Interior Forest of Argentina (with Rosewood & Assai Palm)
These forests found in the extreme north of Misiones on the Iquazú and Paraná river plains were once characterized by an abundance of the two endemic trees Aspidosperma polyneuron Rosewood (Apocynaceae) and Euterpe edulis Assai Palm (Arecaceae), but these have now been reduced to almost relict status. Several local endemic plant taxa occur including Begonia descoleana (Begoniaceae), Cyperus andreanus var. yguazuensis (Cyperaceae), Peperomia misionense (Piperaceae) and Podostemum comatum (Podostemaceae). (To be completed).
Atlantic Interior Forest of Argentina (with Laurel and Guatambu)
Forests characterized by an abundance of Nectandra species (laurels) and the endemic Balfourodendron riedelianum (Rutaceae) are found in the north and central parts of Misiones. In the west they are bordered by the Paraná River, in the east by mountains, and in the south they merge into campos. Parts of this formation occur in the gullies and flood valleys of the Paraná and its tributaries. Here along the banks of the rivers Sapindus saponaria and the endemic or near endemic bamboo Guadua paraquayana (Poaceae) typically occur.
Atlantic Montane Forest of Argentina
Confined to the hills and mountains of central eastern and northeastern parts of Misiones, these forests have much in common with Laurel and Guatambu Forest, but they also have a number of distinctive elements. In particular, they support an abundance of tree ferns of the genera Alsophyla, Dickinsonia and Trichipteris and as a consequence this forest has been called the tree fern district. Other notable plants include Cyperus burkartii and Nothoscordum moconense.
Atlantic Mountain Grasslands
In the highlands of the Itatiaia Range at altitudes between about 2000-2400 m grasslands almost exclusively dominate well-drained areas. They generally grow to about 20 cm in height and typically include Andropogon incanus, Calamagrostis montevidensis, Danthonia montana, Panicum demissum, P. setifolium and the endemic Briza braziliensis and Bromus brachyanthera (Poaceae) together with species of Deschampsia, Festuca and Trachypogon. Isolated woody species may also occur with gnarled trunks and irregular open crowns. They are invariable associated with large boulders presumably as a form of shelter. Among other characteristic species are Clethra laevigata, Roupala gardneriana, R. lucens, R. schwackeana and the endemic Weinmannia discolor (Cunoniaceae). On well-drained slopes particularly where there are low levels of organic matter in the soil dense swards of the endemic dwarf bamboo Chusquea pinifolia (Poaceae) can be found. These can reach heights of 2 m and are almost impenetrable in places. In the flatter, poorly drained areas, grasslands dominated by the tall, endemic grass Cortaderia modesta (Poaceae) usually form the main vegetation type. These tall, bunch-grass formations can range in height from 1-2 m, but individual clumps are often separated by up to 1 m.
Atlantic Mountain Wetlands
In the Itatiaia Mountain Range most of the bogs are dominated by graminoids but usually have a ground layer of Sphagnum moss. In the wettest areas the upper layer is typically dominated by Cladium ensifolium, but when slightly dryer conditions prevail the endemic grass Cortaderia modesta (Poaceae) usually becomes the main species. Associated species include Alstroemeria isabellana, Blechnum schomburgkii, Polystichum quadrangulare and Rhabdocaulon coccineus. The many small swamps are also largely dominated by graminoides particularly of the genera Bulbostylis and Fimbristylis. Sphagnum again dominates the lower stratum but small plants such as Anagallis tenella, Ranunculus montevidensis and the local endemic Hydrocotyle itatiaiaensis (Apiaceae) may also be present. Ponds and lagoons such as Lagoa Bonita, locally referred to as ‘pozzinas’ usually have bottom dwelling species like Isoetes gardneriana and I. martii, while the emergent vegetation typically include Juncus microcephalus and the endemic Juncus ustulatus (Juncaceae) together with several species of Eriocaulaceae. Creeping plants like Lilaeopsis ulei and the local endemic Boopis itatiaiae (Calyceraceae) may also be present. Stream margins are typically dominated by shrubby vegetation in which Azara uruguayensis, Leucothoe rivularis and Myrica hispida usually predominate, while the ground layer of these communities is typically dominated by creeping plants such as Blechnum pennamarina and the endemic Geranium brasiliense (Geraniaceae) and Itatiaia cleistopetala (family?).
Atlantic Mountain Rupicolous Formations
These upland rupicolous or rocky plant formations occur in isolated patches in a number of the Atlantic Serras. On the high plateau of Itatiaia, for example, there are good examples. Here this vegetation can vary considerably over very short distances. Vegetation islands dominated by the local endemic mat-forming species Fernseea itatiaiae (Bromeliaceae) or Pleurostima gounelleana (Velloziaceae) are often in close proximity to wetland species such as Juncus microcephala and the endemic Utricularia reniformis (Lentibulariaceae) and Xyris teres (Xyridaceae), and woody plants like Eupatorium alpestre and the endemic Baccharis stylosa (Asteraceae). On talus slopes, the endemic tank bromeliad Vriesea itatiaiae (Bromeliaceae) is often the main species. These upland areas provide harsh condition for plant life. There is increased exposure to wind and the rocky substratum has low levels of nutrients and water retention. The specialist flora includes species with various adaptations. Some, for example, (mostly members of the Cyperaceae and Velloziaceae in South America) are know as ‘resurrection’ plants in that they are capable of rehydrating desiccated leaves. Other adaptations to the lack of water include atmospheric roots, bulbs and tank-forming leaves. In fact, geophytes, such as the endemic Alstroemeria foliosa (Alstroemeriaceae), Hippeastrum morelianum (Amarylidaceae) and Stevia camporum (Asteraceae) are among the most abundant species with a high frequency in all microhabitats. Therophytes, on the other hand, are rare. The low nutrient budget and short growing season is thought to preclude any rapid establishment and growth. Many other species are endemic to these rocky upland areas and a number of these such as Begonia lanstiakii (Begoniaceae), Briza itatiaiae, Chusquea microphylla (Poaceae), Leiothrix argyroderma, Paepalanthus itatiaiensis (Eriocaulaceae), Salvia itatiaiensis (Lamiaceae), Senecio arygrotrichus (Asteraceae), Viola uleana (Violaceae), Xyris wawrae (Xyridaceae) and the fern Doryopteria feei (Pteridaceae) are confined to Itatiaia. In the Serra das Cabaças part of the Serra do Mantiqueira, cushions of Trilepis lhotzkiana dominate many of the so-called monocotyledonous mats that cover many of the granitic outcrops. Other common associates include Nanuza plicata, Rhynchospora emaciata, Stevia claussenii and the endemic Vellozia candida (Velloziaceae). The most important mat forming plants are species of Bromeliaceae and Velloziaceae, and these mats act as an important substratum for other species to colonize. Other endemic species found here include Baccharis stylosa (Asteraceae) and Sininga magnifica (Gesneriaceae). A defining feature of these communities is their so-called oligarchic structure. In other words, they have few dominant species but support many locally rare species. Similar formations are found on the Gneiss-granite outcrops around Rio de Janeiro (Serra do Mar). Trilepis lhotzkiana is again the most frequent species closely followed by Nanuza plicata, Epidendron elongatum, Pleurostima purpurea, Selaginella convoluta, S. sellowii and the narrow endemic cactus Cephalocereus fluminensis (Cactaceae). Plant endemism is again incredibly high. In fact, certain separated narrow endemics are within seeing distance of each other highlighting the insular character of these outcrops. Among the many narrow endemics are Alcanterea geniculata (Bromelaceae), Anthurium sucrii (Araceae), Manihot leptopoda (Euphorbiaceae), Paliavana racemosa (Gesneraceae), Pleurostima seubertiana (Velloziaceae), Rhipsalis cereoides (Cactaceae), Trilepis ciliatifolia (Cyperaceae) and Vernonia longo-angustata (Asteraceae).
Atlantic Summit Formations
In the summit zones of the Itatiaia Range at altitudes ranging from about 2400-2770 m, even the most favourable, sheltered areas are subject to extremes of wind and temperature. In the most extensive formations, the endemic bamboo Chusquea pinifolia (Poaceae) typically dominates, and can stand up to a height of 1 m. In high, shady, often humid valleys between the highest peaks there are dense shrubby formations growing up to 2 m in height. Typical species here include Griselina ruscifolia, Myrica hispida, Purpurella hospita and the endemic Leandra sulfurea (Melastomataceae). Often mixed with these shrubs is the endemic Fuchsia campos-portoi (Onagraceae). Crevices provide habitat for various low growing herbs and shrubs, a number of which form tight cushions. Typical species include Achyrocline satureoides, Chionolaena glomerata, Lepechinea speciosa and Leuchopholis capitata. On relatively flat areas where the rock has become broken up, such, as on Aqulhas Negras and Prateleiras, there are interesting, compact carpets of Oxalis calva and Zygocactus obtusangulus, which can be up to 10 cm thick.
Atlantic Coastal Sand Dunes
Dunes on the Atlantic coast of Brazil comprise of a large variety of different vegetation zones ranging from foredunes to primary and secondary dune scrub and flooded dune forest.
Foredunes
This zone extends from the upper part of the beach to the top of the foredune crest and may be inundated by seawater during exceptionally high tides. It is therefore dominated by prostrate, largely succulent halophytes. The main species is Blutaporon portulacoides but other frequent species include Alternanthera maritima and Sesuvium portulacastrum.
Primary Dune Zone
This zone can extent from five to a few hundred metres inland and is normally less than 2 m high but can reach heights of up to 100 m on the north-eastern coast where it is frequently mobile and unvegetated. It is typically composed of a creeping psammophytic zone which in places gives way to low closed thicket.
Creeping Psammophyte Primary Dune Zone
This zone is mainly dominated by grasses and sedges such as Mariscus pedunculatus, Panicum racemosum and Sporobolus virginicus but also includes broad-leaved creeping plants like Canavalia rosea, Ipomoea littoralis and Ipomoea pes-caprae. In southern latitudes Spartina ciliata becomes the main species. Plant height rarely exceeds more than about 30 cm (about 1 m in the case of Spartina) and plant cover is usually less than 20%.
Low Closed Thicket Primary Dune Zone
In certain areas such as along the coast of Rio de Janeiro and southern Espirito Santo where there has been less of a human impact the creeping psammophyte zone gives way to dense, dune shrub vegetation. This can reach heights of up to 3 m in the back-dune zone. The most abundant species are Bumelia obtusifolia, Schinus terebinthifolius and the endemic Jacquinia brasiliensis (Theophrastaceae). The zone also includes a number of cacti and bromeliads and the liana Aristolochia macroura.
Swale Zone
Between successive beach ridges are low areas known as swales. These zones are usually less than 300 m but support a huge diversity of species ranging from typical hydrophytes to wet forests. At least four communities have been recognised. These include sedge swamp, periodically flooded forest, permanently flooded forest and Ericaceae open scrub. These all appear to relate to differing ground-water levels
Swamp Sedge Swale
These zones are inundated for most of the year but water depth is usually less than 20 cm. Sedges such as Eleocharis subarticulata, Rhynchospora corymbosa and Rhynchospora holoschoenoides are the dominant species but grasses, such as Panicum pumilum and Paspalum arenarium are common while Tibouchina urceolaris and Xris jupicai are conspicuous along the edges of the flooded zones.
Periodically Flooded Forest Swale
Reaching no more than about 15 m in height these forests are typically dominated by Calophyllum brasiliensis and Symphonia globulifera. The understory usually includes Psychotria carthaginensis and Rheedia brasiliensis while Aechmea pineliana is a frequent herb layer bromeliad. Also frequent is the understory palm Geonoma schottiana.
Permanently Flooded Forest Swale
Dominated by Tabebuia cassinoides, which forms a canopy up to 8 m high, these forests are otherwise relatively poor in species. The few other species include the palm Bactris setosa and small specimens of Symphonia globulifera, while in the understory the fern Acrostichum danaefolium is a characteristic feature.
Ericaceae Open Dune Scrub
These distinctive scrublands are most extensively found on the north-eastern plains of the Rio de Janeiro coast. They are typically found on slightly elevated area where termite hills occur. The characteristic species include Andira legalis, Esterhazya splendida, Gaylussacia brasiliensis, Protium icicariba and the endemic Leucothoe revolute (Ericaceae) and Ocotea notata (Lauraceae). In some areas the area between scrub is covered by a dense stand of the low shrub Marcetia taxifolia. The herbaceous layer typically comprises dense stands of Vriesea procera, and Catasetum discolor is a characteristic ground orchids.
Secondary Dunes
This zone can extend from about 500 m from the ocean inland for several kilometres but in exceptional circumstances can form a zone up to 25 km wide. It eventually grades into Atlantic rainforest. The zone can be resolved into three communities: Myrtaceae closed thicket, Clusia open scrub, and palm open scrub.
Secondary Dunes with Myrtaceae Closed Thicket
Reaching about 4 m in height this zone is dominated by species of the family Myrtaceae with Eugenia and Myrcia being the most important genera. These 2 genera contribute up to about 12 of the species. Other important shrubs are Aspidosperma pyricollum, Cupania emarginata, Esenbeckia rigida, Gomidesia maritime, Melanopsidium nigrum, Myrrhinum atropurpureum, Pouteria psammophila, Rheedia brasiliensis and Zollernia falcate. The herbaceous layer largely comprises Aechmea lingulata, Billbergia amoena and the endemic Quesnelia quesneliana (Bromeliaceae). Bromeliads are reported to be important colonisers of these open dunes.
Secondary Dunes with Clusia Open Scrub
Forming dense often hemispherical thickets interspersed with bare patches these open scrublands can reach heights of up to 4 m. Clusia is the most important genus but species composition varies from site to site. Typical species include Clusia australis, Clusia fluminensis, Clusia lanceolata and the endemic Clusia hilariana (Clusiaceae). Other characteristic shrubs include Cassia australis, Erythroxylum ovalifolium, Eugenia nitida, Gomidesia fenzliana, Manilkara subsericea, Myrica lundiana and Tapirira guianensis. Also characteristic are various climbing plants such as Mandevilla funiformis, Passiflora mucronata, Peixotoa hispidula and the endemic Paullinia weinmanniaefolia (Sapindaceae). The sparse herbaceous layer is characterised by various Anthurium and Polypodium species together with various bromeliads and cacti. Terrestrial bromeliads, such as Aechmea nudicaulis, Vriesea procera and the endemic Neoregelia cruenta, are often found in the borders around thickets. In the sparsely vegetated areas between thickets isolated plants of many species can be found. The most characteristic of these are Allagoptera arenaria, Cassia ramose, Pilosocereus arrabidae, Stachytarpheta schottiana and Stigmaphyllon paralias.
Secondary Dune with Palm Open Scrub
Whenever other forms of secondary dune vegetation is burned on the Rio de Janeiro coast a low scrub community dominated by the palm Allagoptera arenaria often becomes the dominant vegetation. This disclimax can often cover up to 30% of any devastated area. Other important species include Axonopus barbigerus, Byrsonima sericea and Vriesea neoglutinosa.
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