Central American (Yucatan) Tropical Forest
Until recently these forests were thought to be relatively homogenous, but are now known to comprise several communities depending on soil type, terrain and rainfall levels. In the low rainfall areas on relatively shallow calcareous soils in northern parts of Quintana Roo, the forest exclusive to this zone is characterized by Agave angustifolia, Plumeria obtusa and the Yucatan endemic Sebestiana adenophora (family?). These low stature forests rarely exceed about 7 m but typically have a shrub layer up to 2 m and an herb layer up to 1 m. Other common species include Bromelia karatas, Esenbeckia pentaphylla, Eugenia mayana, Rhoeo discolor and the Yucatan endemics Beaucarnea pliabilis (Ruscaceae), Bourreria pulchra (Boraginaceae), Caesalpinia yucatanensis (Fabaceae) and Trichilia minutiflora (Meliaceae). Where rainfall is slightly higher the forests are characterized by Manilkara zapota and the Yucatan endemic palm Coccothrinax readii (Arecaceae). These forests cover many parts of Quintana Roo and can have canopy trees reaching 30 m or so. Other typical species include Bursera simarouba, Laetia thamnia, Malmea depressa, Neea psychotriodes and the Yucatan endemics Hampea trilobata (Malvaceae) and Ottoschulzia pallida (Icacinaceae).
The under storey consists largely of saplings and climbers, but shrubs such as Abutilon permolla, Byttneria aculeata and Malvaviscum arboreus may be encountered together with herbs such as Anthurium schlechtendalii, Lasiacis divaricata and the endemic Olyra yucatana (Poaceae). With increasing rain dense forests typified by Bursera simaruba and the Yucatan endemics Hampea trilobata (Malvaceae) and Metopium brownei (Anacardiaceae) occur. These forests have a central distribution in Quintana Roo. Common understory species include Casearia nitida, Chamaedorea seifrizii, Lonchocarpus rugosus and Randia aculeate, but a special feature of these forests is the rich arboreal flora with herbaceous species such as Ichnanthus lanceolata and Zamia loddigesii, shrubs such as Malvaviscus arboreus and climbers like Arrabidaea floribunda, Passiflora coriacea and the Yucatan endemic Desmonicus quasillarius (family?). In the highest rainfall areas where annual precipitation exceeds 1400 mm the forests normally have three distinct stories. In south-central Quinttana Roo the characteristic species are Caesalpinia gaumeri and Vitex gaumeri. Other typical species include Byrsonima bucidaefolia, Diospyros verae-crucis, Spondias mombin and the Yucatan endemics Acacia gaumeri (Fabaceae) and Alseis yucatanensis (Rubiaceae). Vitex gaumeri is one of the most abundant trees on the peninsula. In the wettest areas in southern Quintana Roo the dense forest have a well-developed vertical structure with canopy trees reaching heights of up to 30 m. Here the most abundant trees are Brosimum alicastrum, Orbignya cohuna and Trichilia glabra. However, the forests throughout the Yucatan are extremely diverse and contain many of the species endemic to the peninsula. In addition to the ones already mentioned these include Acacia cedilloi (Fabaceae), Coccoloba reflexiflora (Polygonaceae), Croton campechianus (Euphorbiaceae), Cryosophila stauracantha (Arecaceae), Diospyros cuneata (Ebenaceae), Erythroxylum bequaertii (Erythroxylaceae), Exothea diphylla (Sapindaceae), Jatropha gaumeri (Euphorbiaceae), Lonchocarpus xuul (Fabaceae), Platymiscium yucatanum (Fabaceae), Sideroxylon foetidissimum subsp. gaumeri (Sapotaceae) and Thouinia paucidentalis (Sapindaceae).
Central American (Costa Rican) Tropical Forest
The La Selva Biological Station located where the foothills of Costa Rica’s central volcanic mountains give way the Caribbean coastal plain has compiled detailed descriptions of these forests. They are characterized by irregular canopies ranging in height from 30-55 m and dominated by the endemic Pentaclethra macroloba (Mimosoidaceae). Palms are a distinctive feature of the sub canopy which are made more conspicuous by the long leaves of Welfia georgii, the high stilt roots of Iriartea deltoidea and Socratea exorrhiza, and the long spines of the endemic Astrocaryum confertum (Arecaceae). Typical dicot species of the sub canopy include Dendropanax arboreus, Dystovomita paniculata and the endemic Protium panamensis, P. pittieri (Burseraceae) and Unoniopsis pittieri (Annonaceae). However, below the sub canopy the under storey is more difficult to define, but a story of small trees ranging in height from 5-15 m can often be distinguished. These include Anaxagorea crassipetala, Cassipourea elliptica, Rinorea deflexiflora, Symphonia globulifera and the endemic Guarea rhopalocarpa (Melicaceae). A story that can be broadly described as a shrub layer usually occurs growing to a height of about 5 m. Palms are again characteristic here with species such as Asterogyne martiana and the endemic, multi-stemmed Geonoma congesta (Arecaceae), while common dicots are Carpotroche platyptera, Mabea occidentalis, Miconia simplex and the endemic Herrania purpurea (Sterculiaceae). Tree seedlings and woody lianas rather than herbaceous plants dominate much of the ground layer, but with occasional perennial herbs. These include Asplundia uncinata, Besleria columneoides, Danaea wendlandii and the endemic Calathea cleistantha (Marantaceae) and Spathiphyllum fulvovirens (Araceae). There are a number of other interesting endemic species found in these forests. Pariana parvispica (Poaceae) is a bambusoid grass endemic to the Caribbean lowland forests of Costa Rica, and represents the northernmost example of this South American genus. Zamia skinneri (Zamiaceae) is an endemic under story cycad with no noticeable difference between the sexes.
Central American (Costa Rican) Swamp and River Forest
Good descriptions of these forests have again been compiled for the La Selva area of Costa Rica. They are mainly confined to wet areas adjacent to rivers and swampy valley bottoms. The swamp forests often flood after heavy rain to a depth of 20-30 cm, but the floodwaters rarely remain for more than a few days. As with Costa Rica’s dryer forests, the endemic Pentaclethra macroloba (Mimosoidaceae) is the dominant tree species, while other more characteristic canopy trees include Luehea seemannii, Otoba novogranatensis, Pachira aquatica and the endemic Carapa nicaraguensis (Meliaceae). A sub canopy can usually be distinguished, but few of these smaller trees, such as the endemic Pithecellobium valerioi (Fabaceae) are confined to swamps. The few more characteristic small trees include the endemic palm Astrocaryum alatum (Arecaceae) and the endemic dicot Chione costaricensis (Rubiaceae). Characteristic shrubs include the endemic Adelia triloba (Euphorbiaceae) and Psychotria chagrensis (Rubiaceae). River bank vegetation also includes a number of characteristic trees. Large trees such as Ficus insipida and Pithecellobium longifolium are typically found arching over rivers such as the Rio Puerta Viejo. Other tree such as Inga marginata, Nectandra reticulata, Posoqueria latifolia and the endemic Cordia lucidula (Boraginaceae) are more or less confined to these riparian habitats. Also characteristic of these riversides are various species more typically found at higher elevations. These include endemic shrubs like Acacia ruddiae (Fabaceae), Cardudovica rotundifolia (Cyclanthaceae), Piper friedrichsthalii (Piperaceae) and the endemic herb Cuphea utriculosa (Lythraceae).
Central American (Costa Rican) Montane Bamboo-Oak Forest
On the Cordillera de Talamanica these forests range in altitude from 2200-3100 m and can be divided into several sub-formations. In general, however, they can be broadly categorized into forests characterized by either the endemic Quercus costaricensis or the endemic Quercus copeyensis (Fagaceae). Quercus costaricensis forest often has five or more vertical layers and can reach heights of up to 30 m. A sub canopy layer is often characterized by the endemic small tree Myrsine pittieri (Myrsinaceae), while other sub canopy trees include Clethra gelida, Drimys grandensis, Oreopanax capitatus, Weinmannia trianae and the endemic Didymopanax pittieri (Araliaceae), Ilex vulcanicola (Aquifoliaceae), Rhamnus oreodendron (Rhamnaceae), Symplocos austin-smithii (Symplocaceae), Vaccinium consanguineum (Ericaceae) and Viburnum costaricanum (Caprifoliaceae). The shrub layer is typically dominated by species of the bamboo genus Chusquea and the endemic shrub Pernettya coriacea (Ericaceae) and can reach heights of 5 m. Below this the herb layer can grow up to 1 m tall and is typically characterized by ferns and terrestrial bromeliads. Finally there is usually a conspicuous bryophyte ground layer often dominated by peat forming Sphagnum species.
In slightly more humid conditions the endemic Quercus copeyensis becomes co-dominant with Q. costaricensis. Other associated trees include Podocarpus oleifolius, Prumnopitys standleyi, Weinmannia pinnata and the endemic Ocotea austinii (Lauraceae). In places the canopy can reach 40 m. The sub canopy ranges from 15-25 m and is again rich in endemic taxa such as Ilex lamprophylla var. discolor (Aquifoliaceae), Miconia pittieri (Melastomataceae), Nectandra salicina (Lauraceae), Prunus annularis (Rosaceae), Zanthoxylum scheryi (Rutaceae) and others. Chusquea bamboos again dominate the shrub layer and there are rich herb and ground layers. In areas where cloud cover becomes an important factor (the cloud belt) Quercus copeyensis usually becomes the dominant canopy tree and epiphytes become a lot more conspicuous often blanketing trees. Here subcanopy trees grow to about 25 m and include endemic species such as Miconia costaricensis (Melastomataceae), Cornus disciflora (Cornaceae), Phoebe pittieri (Lauraceae) together with many of the aforementioned species. In the shrub layer the palm Geonoma hoffmanniana becomes particularly important together with Chusquea bamboos (including C. foliosa and C. patens), several tree ferns, the broad-leaved bamboo Aulonemia viscose, and the endemic shrub Cavendishia crassifolia (Ericaceae). The herb layer is rich in ferns and herbs such as Maianthemum paniculatum and the endemic Begonia udisilvestris (Begoniaceae).
Finally in places were conditions are slightly warmer and possibly drier these forests become co-dominated by the endemic white-barked Quercus copeyensis and the endemic black-barked Quercus seemanni (Fagaceae). The sub-canopy here has many species in common with other sub-formations, but also commonly includes the endemic Oreopanax oerstedianum (Araliaceae). Chusquea is again dominant in the shrub layer together with species such as Fuchsia microphylla and the endemic Psammisia ramiflora (Ericaceae). At ground level herbs and ferns dominate with Adiantum concinnatum, Bomarea tenerum, Pteris muricella and the endemic Anthurium concinnatum (Araceae), Centropogon costaricae (Campanulaceae) and Verbesina oerstediana (Asteraceae) being particularly common.
Central American Mosquitia Pine Savanna
Resembling the pine flats of Florida and Louisiana, these open park-like savannas are mainly characterized by the presence of sedges, particularly Rhynchosporum globosa, and could have been possibly more appropriately called sedge savanna. The most conspicuous tree is Pinus caribaea, while other common woody species are Byrsonima crassifolia and the endemic oak Quercus oleoides (Fagaceae). Rhynchosporum globosa forms ceapitose tussocks while other abundant herbaceous species, often growing between the tussocks, include Paspalum pulchellum, Rhynchosporum barbata, R. chapmannii, Tonina fluviatilis and Utricularia subulata. In slightly dryer areas Bulbostylis paradoxa and Declieuxia fruticosa become more conspicuous. Most of the herbaceous vegetation grows between 2-3 dm tall, but thickets of Paurotis wrightii are common in wet areas. On slightly higher ground shrubs of Davilla kunthii and Quercus oleoides can predominate. Other endemic species include shrubs such Miconia hondurensis (Melastomataceae), Tabernaemontana chrysocarpa (Apocynaceae) and the herb Aciotis rostellata (Melastomataceae).
Central American (Guatemalan) High Elevation Conifer Forests
In the Sierra de los Cuchumatanes and Cadena Volcanica areas these upland conifer forests can be broadly divided in to pine and fir forests. Pine forest dominated by Pinus hartwegii appears to be the main climatic climax community in the altitudinal range of 2800-4000 m. However, these forests are far from homogenous and can be divided into several sub-formations or communities. In places the herb layer is dominated by Agrostis tolucensis and Relbunium microphyllum while less common species include Eryngium scaposum, Plantago tubulosa, Senecio warcewiczii and the endemic Lepechinia schiedeana (Lamiaceae). This type of forests can be found on both flat and undulating terrain but forest with herb layers dominated by Agrostis exserta and Werneria nubigena are confined to undulating areas. Here there is usually a low herb layer with Eryngium cymosa, Gentiana pumilio and Werneria nubigena and tall herb layer with Agrostis exserta and Helenium integrifolium. On steep the slopes of Cadena Volcanica, open, monodominant stands of Pinus hartwegia occur that can reach heights of 20 m or more. Here the under storey if largely composed of a bunch grass layer with Calamagrostis vulcanica and Lupinus montanus and a ground layer dominated by Lachemilla vulcanica. Other species include Castilleja integrifolia, Gnaphalium salicifolium and the endemic Senecio oerstedianus (Asteraceae).
On less steep slopes Pinus hartwegii is joined by the sporadic occurrence of trees such as Alnus firmifolia and the endemic Buddleja magalocephala (Scrophulariaceae) and a shrub layer comprising Holodiscus argenteus, Rubus trilobata and the endemic Juniperus standleyi (Cupressaceae), which can reach heights of up to 6 m. Most of the pine trees in this zone appear to be infested by the hemiparasitic plant Arceuthobium globosum (Loranthaceae). Other endemic species associated with these conifer forests include Cerastium guatemalense (Caryophyllaceae), Cirsium skutchii, Hieracium guatemalensis, Eupatoria nubivagum, Senecio nubivalus (Asteraceae), Cardamine eremita (Brassicaceae) and Poa venosa (Poaceae).
Fir forests dominted by the endemic Abies guatemalensis (Pinaceae) are characteristic of very steep slopes ranging in elevation from about 2800-3400 m. These are dense forests that grow to a height of 30 m or so. Other characteristic species include Sabazia pinetiorum. A distinct shrub layer with species such as Rubus trilobus, Senecio callosus and Tetragyron orizabense is usually present but a conspicuous herb layer is largely absent. Other characteristic species include Cheilanthes chaerophylla, Salvia excelsa, Trifolium amable and the endemic Arenaria guatemalensis (Caryophyllaceae).
Central American Tropical Dry Forest
Originally these forests stretched along the Pacific coast of Central America from southwest Mexico (southern Chiapes) through Guatemala, El Salvador, Honduras, and Nicaragua to northwestern Costa Rica, in a zone where there is a prolonged dry season, but today these forests are highly fragmented. In altitude they range from sea level to about 800 m and are usually low stature, semi-deciduous forests with no more than two vertical tree stories. Certain trees can, however, reach heights of up to 30 m. In southern sectors the common trees include Annona holosericea, Bursera simoruba, Casearia arguata, Eugenia salmensis, Guazuma ulmifolia, Jacquinia pungens, Tabebuia chrysantha and a variety of endemic and near endemic taxa such as Bombacopsis quinatum (Bombacaceae), Brosimum costaricensis (Moraceae), Dalbergia retusa (Fabaceae), Enterolobium cyclocarpum (Fabaceae) and Jacquinia macrocarpa (Theophrastaceae). Most of the trees belong to the Leguminosae superfamily of Fabaceae and many have associations with nitrogen fixing bacteria or have symbiotic relationships with various species of ant. The presence of Bombacopsis quinatum is of considerable biogeographical interest since it represents a family (Bombacaceae), which includes the baobabs of Africa and Australia. These are said to be descendents of Gondwanan ancestors and the presence of Bombacopsis quinatum in the Neotropics is quoted as evidence of continental drift. Bursera simoruba (naked indian) is of interest for its ability to photosynthesis through its shiny green bark. The under storey may include various shrubs such as the endemic Cupania guatemalensis (Sapindaceae), Machaerium biovulatum (Fabaceae) and Miconia argentea (Melastomataceae), clumps of palms (mainly species of Bactris) and occasional terrestrial bromeliads. Epiphytes are sparsely distributed and not conspicuous, but the crowns of trees are often lightly festooned with orchids, bromeliads, mosses, lichens, and there are occasional lianas.
Central American (Yucatan Peninsula) Coastal Dunes
Coastal dunes on the Yucatan Peninsula can be broadly divided into beach, embryo dunes, foredunes, wet and humid slacks, humid sheltered dunes and dry dunes. They tend to be sheltered apart from a few hurracanes and their coarse sand prevents much mobility - there are no blowouts, for example. Overall they support about 31 dune community types. In terms of their physiognomy and floristics they are similar to the dunes of the Caribbean islands.
Beach (Pioneer Zone)
Salt spray and wind are the main factors here but there is little sand movement and the vegetation tends to be monospecific. Typical species include Cakile edentula, Cakile lanceolata and Suaeda linearis.
Embryo Dunes
The main species here are Canavalia rosea, Euphorbia buxifolia, Ipomoea pes-caprae, Sesuvium portulacastrum and Sporobolus virginicus. All are classed as nitrophiles able to tolerate the high nutrient enrichment resulting from stranded material and sea spray.
Foredunes
Most of the foredune vegetation consists of woody species standing up to 4 m high in places. They are all tolerant of salt spray and sand deposition and characteristically have succulent leaves. Depending on location the dominant species may include Croton punctatus, Euphorbia buxifolia, Scaevola plumieri, Suriana maritima or Tournefortia gnaphalodes.
Humid and Wet Slacks
These are not typical slacks but more accurately described as flat humid plains flooded during summer. They also receive more salt spray than typical slacks and most species are salt tolerant. In fact, the herbaceous flora has much in common with the saltmarshes of North America. Typical species include Ambrosia hispida, Conocarpus erectus, Eleocharis geniculata, Flaveria linearis, Gossypium hirsutum, Hymenocallis americana, Hymenocallis caribaea, Iresina stricta, Lippia nodiflora, Lycium carolinianum, Maytenus phylantoides, Melanthera nivea, Opuntia stricta, Scaevola plumieri, Sesuvium portulacastrum, Suaeda linearis, Suriana maritima and Waltheria indica.
Fixed Dunes
In humid areas thickets up to 4 m tall can develop and usually have a well developed vertical structure. These typically have three layers comprising small trees and palms, shrubs and an inconspicuous herb layer and all have some tolerance to saline conditions. Depending on location the dominant species may include the shrubs Chrysobalanus icaco or Ernodea littoralis, the small tree Coccoloba uvifera or the palms Pseudophoenix sargentii or Thrinax radiata. The endemic Coccothrinax readii and poisonous Metopium brownei is often found in association with Thrinax radiata.
In dry areas thickets dominated by succulent spiny species develop with few palms but herbs are more abundant. Depending on location the characteristic species may include the shrubs Bravaisia tubiflora, Brumelia retusa, Caesalpinia vesicaria or Pithecellobium keyense or the small tree Cordia sebestena.
In open dry areas on ridges independent of ground water levels the characteristic species are either Agave angustifolia or endemic shrub Enriquebeltrania crenatifolia (Rhamnaceae). However the former may be derived from planted populations since it was previously cultivated for ‘sisal’ leaf export. Frequent associates in the stands of Enriquebeltrania crenatifolia include the herb Justicia carthaginensis and succulent, thorny shrubs such as Acanthocereus pentagonus while Cyrtopodium punctatum and Schomburkia tibiscinis are common field layer orchids.
Central American (Gulf of Mexico) Coastal Dunes
The Gulf coast can be broadly divided into beach, embryo dunes, foredunes, blowouts, active dunes, wet and humid dune slacks and dry dunes. Overall they supports some 58 dune community types.
Beach (Pioneer Zone)
This zone, which is often subject to damaging hurricanes and inundation from sea water and flooding from nearby rivers, is dominated exlusively by annual species. These help bind sand but do not form embryo dunes. Species composition and dominance varies but typical dominants includes Amaranthus greggii, Cakile edentula, Cakile geniculata, Ipomoea stolonifera, Okenia hypogaea and Sporobolus virginicus.
Embryo Dunes and Foredunes
Species composition of embryo dunes and foredunes are similar but they all constitute species that can tolerate large amounts of sand accretion and salt spray, and share an ability to trap sand. Some of the more dominant species include Bidens pilosa, Canavalia rosea, Croton punctatus, Heliotropium curassavicum, Ipomoea pes-caprae, Ipomoea stolonifera, Oenothera drummondii, Panicum geminatum, Sesuvium portulacastrum, Sporobolus virginicus, Uniola paniculata and the endemic Palafoxia lindenii (Asteraceae). The latter is a tall shrub that can form monospecific stands in places.
Blowout and Active Dunes
These are area of high mobility and low organic content. Typical species include Acacia cornigera, Andropogon scoparius var. littoralis, Chamaecrista chamaecristoides and Commelina erecta. The small shrub Chamaecrista chamaecristoides is considered to be endemic to the Gulf of Mexico. Nevetheless, it is one of the most widespread species in the Gulf and one of the most important dune fixers.
Humid and Wet Slacks
These develop where the ground water is close to the surface. Species composition and dominance varies, but some of the more typical dominants include Bidens pilosa, Borrichia frutescens, Chamaesyce dioica, Conocarpus erecta, Cyperus articulatus, Eleocharis cellulosa, Eleocharis geniculata, Erigeron myrionactis, Fimbristylis castanea, Fimbristylis spathacea, Hydrocotyle bonariensis, Lippia nodiflora and Panicum repens.
Fixed Dunes
In the Gulf these more stable dune areas are far more complex than in temperate dune systems and can be divided into forb dominated communities, grass dominated communities, low shrub dominated communities, spiny shrub dominated communities and high thickets.
Forb Dominated Fixed Dunes
Depending on location the dominant species may include either Macroptilium atropurpureum or Schrankia quadrivalvis.
Grass Dominated Fixed Dunes
Depending on location the dominant species may include Aristida adscensionis, Aristida roemeriana, Aristida curvifolia, Bouteloua repens, Eustachys petraea, Sporobolus jacquemonti or the endemic Trachypogon gouini.
Low Shrub Dominated Fixed Dunes
Depending on location the dominant species may include Ambrosia artemisiifolia, Pectis saturejoides, Stemodia tomentosa, Palafoxia texana var. robusta, Tamonea curassavica or Waltheria indica.
Spiny Shrub Dominated Fixed Dunes
These are often associated with sheltered areas of beach-ridge systems. Depending on location the dominant species may include Acacia macracantha, Acacia farnesiana, Caesalpinia bonduc, Opuntia stricta var. dillenii or Randia laetevirens. The latter is the most widespread of the spiny shrubs forming thickets of between 1-3 m tall.
High Thicket Dominated Fixed Dunes
Depending on location the dominant species may include Chrysobalanus icaco, Coccoloba barbadensis, Crossopetalum uragoga, Dalea scandens, Diphysa robinioides, Eupatorium odoratum, Hibiscus pernabuscensis, Lantana camara, Nectandra coriacea, Psidium guajava, Schaefferia frutescens or Verbesina persicifolia.
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