Formations in the Chocóan BioProvince

Peter Martin Rhind

Chocóan Upland Rain Forest (Bajo Calima) These forests occur on low hills composed of Tertiary sediments. The soils are very low in nutrients and possibly have the lowest phosphorus levels of any known Neotropical forest. At canopy level the palm Jessenia batava appears to be the most common , but other palms such as Socratea exorrhiza and species of Wettinia are also common and, in fact, these forests appear to support more palm species than any other part of the World. Other dominant canopy include Eschweilera panamensis (), Manilkara bidentata (Sapotaceae) and lehmannii (), while other important trees are utile (), Miconia punctata (Melastomataceae), Micropholis crotonoides (Sapotaceae), Pouteria buenaventurensis (Sapotaceae), Tovomita weddelliana (Clusiaceae), Welfia georgii () and Wettinia quinaria (Arecaceae). However, the overstory or canopy of these forests is extremely rich with up to 258 species recorded in just one hectare. Important families in terms of species-richness include Annonaceae, Arecaceae, Hypericaceae, , Melastromataceae, and Sapotaceae. These forests are also especially important for the Burseraceae in terms of species-richness, endemism and ecological dominance, and for the prevalence of especially the Quararibea. In fact, trans-Andean forests appear to be the centre of diversity for this genus. The under storey is variable but typically includes Mabea occidentalis (), Macrolobium archerii (Fabaceae), Miconia centronoides (Melastomataceae), Pausandra trianae (Euphorbiaceae), and Tetrorchidium ochroleucum (Euphorbiaceae), and can, in fact, support up to three times as many trees as the over storey. The most important shrub families are Melostomataceae, Piperaceae, Rubiaceae and Solanaceae. There is a very high diversity of climbers although most of these are hemi epiphytes, including species of Cavandisha, Clusia, Drymonia, , Marcgravia, Norantea, Schradera and Topobea, rather than free- climbing . Normal epiphytes are also well represented particularly by the , but also include many bromeliads, cycanths, gesneriads, orchids, and peperomias. Among the terrestrial herbs Acanthaceae, Commelinaceae and Gesneriaceae are well represented.

Chocóan Coastal Wet Forest On the basaltic rocks of Cape Corrientes, where there is a short dry season between January and March, the closed canopy wet forest reaches heights of up to 45 m and has emergents extending up to 59 m. The emergent species include (Moraceae), amygdaliferum () and Ceiba aff. pentandra (), with many of these tall trees having large buttress roots. The most abundant canopy species are Otoba latialata and Otoba novogranatensis (Myristicaceae), (Arecaceae), Welfia regia (Arecaceae) and Wettinia quinaria (Arecaceae). Here the most abundant families are Arecaceae, Clusiaceae, Fabaceae, Moraceae, Myristicaceae and Sapotaceae. The largely open under storey is mainly composed of smaller versions of the species mentioned above but also includes Conostegia setosa (Melastomataceae). Arecaceae is the most abundant family followed by Fabaceae, Melastomaceae, Moraceae and Myristicaceae. In terms of species-riches, 124 species with basil diameters greater than 10 cm were recorded in a one-hectare plot and these comprised 86 genera and 40 families. This forest like other Chocó forests was found to be notable for its abundance of arboreal palms and possibly has a higher number of palm species than any other Neotropical forest. Also in line with other Chocó forests the hemi epiphytes are more prevalent than lianas.

Chocóan (Tumbesian) Acacia Thorn-Forest Thorn-forest characterized by Acacia species such as A. macracantha and A. tortuosa can replace other natural forests when they have been heavily degraded, but they are also thought to occur naturally on the lower slopes of the Tumbesian zone. These normally form open stands between 5-10 m tall and are largely . Common trees include Bursera graveolens (Burseraceae), Caesalpinia corymbosa (Fabaceae), Capparis mollis (), Muntingia calabura (Muntingiaceae), and where there is some groundwater influence Celtis iguanea (Cannabaceae) and Zizyphus piurensis (Rhamnaceae). Typical shrubs are Capparis cordata (Capparaceae), Cercidium praecox (Fabaceae), Cereus macrostilbas (Cactaceae), Cryptocarpus pyriformis (Nyctaginaceae), Encelia canescens (), Galvezia limensis (Plantaginaceae), Grabowskia boerhaviifolia (Solanaceae), Isocarpus microcephala (family?), Maytenus orbicularis (), Mimosa acantholoba (Fabaceae), Parkinisonia aculeata (Fabaceae) and Vallesia dichotoma (Apocynaceae). The deciduous herb layer develops into a dense, lush and fairly high structure during the rainy season. The main species include grasses and forbs such as Anthephora hermaphrodita (Poaceae), Aristida adscensionensis (Poaceae), Bouteloua disticha (Poaceae), Chloris virgata (Poaceae), Heliotropium angiospermus (Boraginaceae), Luffa operculata (Cucurbitaceae), Nicandra physaloides (Solanaceae), trichotoma (Asteraceae) and Sicyos chaetocephalus (Cucurbitaceae).

Chocóan (Tumbesian) Ceiba trichistandra Forest This largely lowland forest ranges from mainly deciduous to semi-evergreen. The bulbous character of the evergreen, dominant Ceiba trichistandra (Malvaceae) make these forests very distinctive and in the more deciduous stands spiny red-barked Erythrina velutina (Fabaceae) adds to the distinctiveness. Undisturbed forest usually has a closed canopy that can reach heights of 25 m but with certain trees reaching 35 m. The other main canopy trees in the mainly deciduous forests of Tambo Negro include Cochlospermum vitifolium (Bixaceae), Eriotheca ruizii (Malvaceae), Erythrina velutina (Fabaceae), Hura crepitans (Euphorbiaceae) and Tabebuia chrysantha (Bignoniaceae). A sub canopy of small trees and shrubs includes Achatocarpus nigricans (Achatocarpaceae), Albizia multiflora (Fabaceae), Armatocereus cartwrightianus (Cactaceae), Capparis heterophylla (Capparaceae), Carica parviflora (Caricaceae), Croton fraseri (Euphorbiaceae), Ficus americana (Moraceae), Guazuma ulmifolia (Sterculiaceae), Phyllanthus anisolobus (), Pithecellobium excelsum (Fabaceae), Rauvolfia tetraphylla (Apocynaceae), Senna atomaria (Fabaceae), Trichilia hirta (), Triplaris cumingiana (Polygonaceae) and Urera baccifera (). The herb layer is draught deciduous and only conspicuous during the rainy season but contains few grasses. Annuals and geophytes like Adiantum raddianum, Brachiaria fasciculata, Coursetia caribaea and Talinum paniculatum represent much of the ground cover. The few grasses may include Andropogon bicornis, Aristida adscensionensis, Chloris virgata and Pennisetum occidentale. Cacti, orchids, and bromeliads like Guzmania monostachya, Pitcairnia prolifera, Vriesea espinosae and various species of Tillandra, dominate the epiphytes. Other epiphytes include Rhipsalis micrantha and Trichocentrum tigrinum, but overall the epiphytic flora although abundant is not particularly rich in species. Climbers are conspicuous and seem to include a high proportion of cucurbit taxa such as Apodanthera, Momordica and Sicyos.

Chocóan (Tumbesian) Low Montane Cloud Forest Situated on the lower seaward slopes at altitudes ranging from 500-1500 m these forests are constantly subject to very humid conditions because of condensation from frequent cloud cover and is little affected by the dry season. In the Buenaventura area these often open canopy forests can reach heights of 35 m and have emergents extending up to 45 m. Depending on location some of the characteristic tree species include Centrolobium paraense, Podandrogyne brevipedunculata, Matisia cordata, Ochroma pyramidale, Mabea occidentalis, Muntingia calabra, Rheedia edulis, Symphonia globulifera, Spondias purpurea, Tabebuia guayacan, Vemonia baccharoides, Banara guianensis, Conostegia cuatrecasii, Ossaea boeckii, Cedrela odorata, Ficus gigantorice, Myrica pubescens, Bocconia integrifolia, Coccoloba obovata, Oreocalyx grandiflora, Turpinia occidentalis, americanus, Urera caracasana and various species of Miconia. The open conditions allow sufficient sub canopy light for dense middle and under storey formations, and as expected the epiphytic flora is well developed. In fact, dense mats of epiphytes cover much of the canopy branches. These include numerous lichens, bryophytes, ferns, orchids, and species of Araceae and bromeliads. The latter include a large number of Guzmania species such as G. angustifolia, G. garciaensis and G. hitchcockiana which here are much more prevalent than Tillandsia species which tend to be more numerous in less humid conditions. The few species included T. acosta-solisii, T. narthecioides, T. pseudotetrantha and T. venusta.

Chocóan (Darien) Tropical Moist Forest These forests are notable for the size, number and height of the trees. The canopy can reach heights of 40 m and further characterized by the giant emergents like Cavanillesia platanifolia and Ceiba pentandra. Both of these are deciduous and their naked branches can be a striking feature from the air. Other important canopy trees include Anacardium excelsum, Bombacopsis sessilis, Enterolobium cyclocarpum, Licania hypoleuca, Platypodium elegans, Pseudobombax septenatum, apetala, Terminalia amazonica, Tetragastris panamensis, Vitex masoniana and the Panamanian endemics Copaifera aromatica (Fabaceae), Lecythis tuyrana (Lecythidaceae) and Gyranthera dariensis (Bambacaceae). At least 49 canopy trees have been identified. However, the canopy is often broken into several strata, with the Panamanian endemic palm Sabal allenii (Arecaceae) being one of the most abundant sub canopy species, but overall the sub canopy is the richest stratum in terms of numbers of species with at least 157 recorded. Other common sub canopy trees include Allophylus occidentalis, Calycophyllum candidissimum, guidonia, Heisteria longipes, Luehea seemannii, Mouriri parviflora, Neea amplifolia, Ouratea lucens, Quararibea asterolepis, Randia armata, Scheelia zonensis, racemosa, Xylopia aromatica and the Panamanian endemics Annona spaguei (Annonaceae), longipes (Moraceae), Ceiba rosea (Bambacaceae), Centrolobium yavizanum (Fabaceae), Eschweilera woodsoniana (Lecythidaceae), Guapira standleyiana (Nyctaginaceae), mucuna (Fabaceae), Machaerium chambersii (Fabaceae), Pentagonia brachyotis (Rubiaceae), Piper pinoganense (Piperaceae), Protium panamensis (Burseraceae), Rinorea brachythrix () and Swartzia darienensis (Fabaceae). The shrub layer is the second richest stratum with at least 56 species recorded. Some of the more common species include Faramea luteovirens, Mabea occidentalis and the Panamanian endemics Miconia rufostellulata (Melostomataceae), Piper darienense (Piperaceae) and Steriphoma macranthum (Capparaceae). are also very abundant. Common woody vines include Connarus panamensis, Doliocarpus dentatus, Maripa panamensis, Passiflora vitifolia, Rourea glabra, Tetracera volubilis, Wulffia baccata and the Panamanian endemics Anthodon panamense (), Bauhinia eucomosa (Fabaceae) and Desmoncus isthemensis (Arecaceae). Herbaceous vines are typically less common but include Bonamia maripoides, Centrosema plumieri, Dioclea guianensis, Eupatorium macrophyllum, Gurania coccinea, Mendoncia lindavii and Rhynchosia pyramidalis. Epiphytes are occasionally encountered with Tillandsia kegelliana being the most abundant species. Other relatively common species include Aechmea tillandsioides, crassinervium, Condonanthe crassifolia, Drymonia serrulata, Hylocereus monacanthus, Philodendron tripartitum, Wittia panemensis and the Panamanian endemic Topobea pluvialis (Melastomataceae). At ground level at least 35 under storey herbs and 30 palmitas have been recorded. Typical herbs include Aneilema geniculatum, Begonia filipes, Chrysothermis friedrichstaliana, Dichorisandra hexandra, Elephantopus rosea, Petiveria alliacea, Rhynchospora cephalotes, Scleria melaleuca and the Panamanian endemic Peperomia brevipeduncula (Piperaceae). The most common palmitas are Heliconia psittacorum, H. subulata, Renealmia karatas and Xiphidium caeruleum, while other less common species include Aechmea magdalenae, Bromelia karatas, Calathea allouia, Carludovica pittieri, Corozo oleifera, Costus nutans, Ischnosiphon leucophaeus, Renealmia aromatica, Stromanthe lutea and the Panamanian endemics Dieffenbachia pittieri (Araceae) and Phytelephas seemannii (Arecaceae).

Chocóan (Darien) Premontane Wet Forest Located largely in the San Blas cordillera at elevations ranging from 250-600 m these forests characteristically have a dense, evenly closed canopy with many sub canopy palms. The canopy rarely exceeds 30 m and most of the trees are relatively slender although many have buttress or stilt roots. Among the most abundant canopy trees are Anacardium excelsum, Bombacopsis quinata, B. sessilis, Brosimum quianense, Ceiba pentandra, Myroxylon balsamum and Oleiocarpon panamense while listed among the less common species are various Panamanian endemics such as Couroupita magnifica, Lecythis tuyrana, L. elata (Lecythidaceae), Mimusops dariensis (Sapotaceae) and Platymisicum dariense (Fabaceae). A number of canopy species are draught deciduous. The sub canopy is relatively open and in addition to many palms also includes numerous other trees and tree ferns. Some of the most common are Brownia rosa-de-monte, Cespedesia macrophylla, Chomelia spinosa, Conostegia xalapensis, Gustavia superba, Heisteria longipes, cornata, Luechea seemannii, Miconia borealis, Oenocarpus panamanus, Ouratea lucens, Pentagonia macrophylla, Pogonopus speciosus, Posoqueria latifolia, Pourouma scabina, Siparuna pauciflora, Stemmadenia grandiflora, and Xylopia frutescens. The subcanopy also include a number of Panamanian endemics such as Cecropia longipes (Moraceae), Ceiba rosea (Bombacaceae), Gloeospermum portobellensis (Violaceae), Guapira standleyiana (Nyctaginaceae), Guatteria panamensis (Fabaceae), Inga saffordiana (Fabaceae), Macrolobium pittieri (Ochnaceae), Eschweilera panamensis (Lecythidaceae) and Pentagonia brachyotis (Rubiaceae). The dense canopy allows little light to penetrate and so the under storey tends to be fairy open. Nevertheless there are a number of terrestrial herbs. The most abundant of these are Aphelandra sindairiana, Clavija mezii, Clidemia dentata, Coccoloba coronata, Conostegia bracteata, Faramea luteovirens, Hasseltia floribunda, Henrietella fascicularis, Herrania purpurea, Leandra dichotoma, Mabea occidentalis, Palicourea guinensis, Picramnia antidesma, Pothomorphe petata, Psychotria involucrata and a variety of Miconia, Piper and Psychotria species. Several Panamanian endemics also occur including Coccoloba manzanillensis (Polygonaceae), Mollinedia darienensis (Monimiaceae), Piper lucigaudens (Piperaceae) and Topobea pluvialis (Melastomataceae). Lianas, vines and epiphytes are also common. Two of the most abundant woody vines are Drymonia spectabilis and Passiflora vitifolia while some of the less common include Strychnos dariensis (Loganiaceae), which is probably endemic to . Herbaceous vines seem less frequent but include the Panamanian endemic Clidemia oblonga (melastomataceae). Epiphytes are mostly found in the canopy or on the boles of trees and typically include Begonia guaduensis, Columnea consanguinea, Drymonia serrulata, Epidendrum crassilabium, Guzmania minor, Monstera adansonii, Philodendron piperoides, Sphyrospermum buxifolium, Tillandsia crispa and a variety of Anthurium species. At ground level the most abundant herbs include Coccocypselum herbaceum and Spigelia anthelmia while among the less common species are the Panamanian endemics Begonia garagarana (Begoniaceae) and Peperomia urocarpoides (Piperaceae).

Chocóan (Magdalena) Moist Forest These forests, situated in the central Magdalena Valley area and extending to the Uraba and Chocó, can be described as dense, high jungle dominated by large trees such as Anacardium excelsum, Caryniana pyriformis, Caryocar amygdaliferum, Cedrela odorata, Ceiba pentandra, Cordia gerascanthus, Hymenea courbaril, Miroxylon balsamum, Ochroma lagopus, Schyzolobium parahybum, Tabebuia rosea and many more. Several forests strata may be distinguished, and like Darien forests, palms dominate the sub canopy. Vines and epiphytes are also well represented with some 150 epiphytic orchid species recorded. Others include local endemic heliconias like Heliconia laxa, H. lentiginosa, H. rigida and H. sanctae-theresae (Heliconiaceae). The undergrowth includes a rich diversity of species including a number of stunning orchids such as Cycnoches chlorochilon, Peristeria elata, Psychopsis papilio and the giant, local endemic warscewiczii ().

Chocóan (Magdalena) Dry Forest Located around the upper Magdalena River area in a flat inter-Andean valley these dry forests grow to heights of no more than about 15 m, and many of the trees have a characteristic umbrella shape. Typical trees and bushes include Acacia farnesiana, Bulnesia carrapo, Capparis odoratissima, Fagara pterota, tinctoria, Prakinsonia aculeata, Prosopis juliflora and the local endemic Pithecellobium bogotense (Fabaceae). Intermixed with the woody vegetation are various characteristic cacti species like Acanthocereus humilis, Pilosocereus colombianus and Stenocereus griseus many of which have very limited geographical distributions. Other local endemics include Amaria petiolata (family?), Cattleya trianaei (Orchidaceae) the national of Colombia, and Steriphoma colombiana (Capparaceae).

Chocóan (Magdalena) Montane Forest Situated on the inner slopes of the Eastern and Central cordilleras above the Magdalena River these wet forests are often enveloped in cloud and at elevations above 1800 m could be described as cloud forest. The most important trees are Anacardium excelsum, Aniba perutilis, Cedrela odorata, Cordia alliodora, Decussocarpus rospigliossi, Podocarpus oleifolius, Quercus humboldtii, Tabebuia rosea and Vochysia ferruginea. Palms are also an important feature with species such as alpinum, C. parvifrons, C. quindivense, C. vogelianum, Dictyocaryum lamarckianum and the local endemic species Ceroxylon sasaimae (Arecaceae). Other local endemics include crispum (Orchidaceae) and various heliconias such as Heliconia abaloi, H. estiletioides, H. huilensis, H. laxa, H. mutisiana, H. oleosa and H. reptans (Heliconiaceae).

Further information required.

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