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D099 Bursera simaruba - Enterolobium cyclocarpum - Ceiba aesculifolia Dry Forest & Woodland Division

Type Concept Sentence: This division includes seasonally drought-deciduous to semi-deciduous tropical forests distributed from sea level up to 1400 m elevation throughout the Caribbean Basin, southern Gulf of Mexico, and in the Pacific Basin from the Gulf of California through Panama.

Common (Translated Scientific) Name: Gumbo Limbo - Elephant Ear - Pochote Dry Forest & Woodland Division

Colloquial Name: Caribbean-Mesoamerican Dry Forest & Woodland

Hierarchy Level:  Division

Type Concept: This division includes tropical forests characterized by a dry season of several months, and found close to the coast or in interior valleys throughout Mexico, Central America, and the Caribbean Basin. Forests tend to have low stature (5-15 m height), high density of small and medium-sized trees, and typically have a single layer, with somewhat open canopies and no emergent trees. The herb layer may be poorly developed or completely lacking. Floristic diversity is low compared to more humid tropical forests, and a few species may dominate the stands. The following list of species are among those that are diagnostic for this division: Astronium graveolens, Bombacopsis quinata, Bursera simaruba, Bursera excelsa, Bursera fagaroides var. elongata, Bursera instabilis, Bursera morelensis, Calycophyllum candidissimum, Ceiba aesculifolia, Ceiba acuminata, Celtis ehrenbergiana, Coccoloba diversifolia, Conzattia sericea, Enterolobium cyclocarpum, Erythroxylum areolatum, Eugenia axillaris, Exostema caribaeum, Exothea paniculata, Guazuma ulmifolia, Guettarda krugii, Guaiacum sanctum, Guapira obtusata, Gymnanthes lucida, Hypelate trifoliata, Jarilla heterophylla, Krugiodendron ferreum, Maclura tinctoria, Metopium toxiferum, Plumeria obtusa, Sideroxylon foetidissimum, Sideroxylon salicifolium, Martinella obovata, and Tabebuia ochracea. Variation in the dry season period, topography, and substrate determine the specific characteristics of the vegetation types in this division since all of these have great importance in dry forests as determinants of variation in water availability. Seasonal climates with 600-1500 mm rainfall per year and the dry season of two to several months are characteristic. These forests most typically occur in sandy lowlands, littoral or sub-littoral flatlands with rock outcrops and higher terraces facing the sea, karstic flats, towerlike karstic hills, humic carbonate soils, and shallow red ferrallitic soils. These forests tend to occur adjacent and/transitional to more humid submontane tropical forests, or towards more xeric tropical thornscrub vegetation types.

Diagnostic Characteristics: Seasonally deciduous and semi-deciduous forest occurring in tropical latitudes of Mexico, the Gulf of Mexico, Central America, and Caribbean Basin. Species diagnostic for this division include Astronium graveolens, Bombacopsis quinata, Bursera simaruba, Bursera excelsa, Bursera fagaroides var. elongata, Bursera instabilis, Bursera morelensis, Calycophyllum candidissimum, Ceiba aesculifolia, Ceiba acuminata, Celtis ehrenbergiana, Coccoloba diversifolia, Conzattia sericea, Enterolobium cyclocarpum, Erythroxylum areolatum, Eugenia axillaris, Exostema caribaeum, Exothea paniculata, Guazuma ulmifolia, Guettarda krugii, Guaiacum sanctum, Guapira obtusata, Gymnanthes lucida, Hypelate trifoliata, Jarilla heterophylla, Krugiodendron ferreum, Maclura tinctoria, Martinella obovata, Metopium toxiferum, Plumeria obtusa, Sideroxylon foetidissimum, Sideroxylon salicifolium, and Tabebuia ochracea.

Rationale for Nominal Species or Physiognomic Features: No Data Available

Classification Comments: Owing to extensive land conversion and degradation (e.g., from livestock grazing), there remains much ambiguity and unevenness in documentation of this division. More work is needed to fully differentiate this type from slightly more humid and evergreen-dominant, more xeric thornscrub, and tropical Quercus-dominated woodland vegetation types.

Similar NVC Types: No Data Available
note: No Data Available
Physiognomy and Structure: These forests have low stature (5-15 m; extremes up to 25 m height), high density of small and medium-sized trees, and typically have single-layer, somewhat open canopies with no emergent trees. Forests in the humid end of the range are semi-deciduous and taller (up to 20-25 m high), while the forests that grow in areas of lower rainfall are nearly 100% deciduous and generally have a lower height and more open canopy below 15 m. The herb layer may be poorly developed or completely lacking. The dry forests of south Florida and the Caribbean tend to have a greater density of individual stems and shorter canopy heights than tropical dry forests in the mainland Neotropics. Density varies from 14,000 stems >2.5 cm/ha in Puerto Rico dry forest to 4600 stems/ha in Florida (Gillespie 2006, Lugo et al. 2006). Drought-deciduousness is the principal adaptive mode of tropical dry deciduous forests, and at the dry extremes, small evergreen trees are important (Lugo et al. 2006). In the Caribbean, the canopy is seasonally open; there are some truly deciduous species in these forests, with early-successional forests dominated by broadleaf semi-deciduous species and late-successional forests dominated by broadleaf evergreen species. Also in the Caribbean, there tends to be a notable lack of lianas compared with dry forests in the mainland Neotropics (Gillespie 2006). Despite relatively low species diversity, high lifeform diversity is common and is accompanied by diversity in plant habit, leaf size and structure, drought tolerance and growth seasonality; this diversity is attributed to habitat heterogeneity coupled with strong rainfall seasonality (Lugo et al. 1978, Medina 1995, as cited in Lugo et al. 2006). Epiphytic orchid and bromeliad species are often found in areas where frosts or anthropogenic disturbances have not occurred in a long time. During the dry season there is an accumulation of litter because much vegetation is deciduous (Holbrook et al. 1995) and sunlight penetrates to the forest floor which reduces the rate of decomposition by decreasing the relative soil moisture (Pennington et al. 2006). The floral and fruiting phenology is highly seasonal and many species flower synchronously during the transition between the dry season and the rainy season when the trees are still leafless (Bullock et al. 1995). Where these forests occur on karst, they are characterized by trees of small diameter, high tree density, and leaf scleromorphy. These stands have a tendency to show signs of being exposed to frequent drought conditions. This is probably due to the rapid rate of runoff and infiltration of rainwater, low water storage in shallow soils, and high sunlight. But this varies depending on the landscape position and the substrate. For example, at the base of limestone "mogotes" or towerlike "haystack mountains" the vegetation may quickly transition to mesic, closed canopy of evergreen species 25-30 m tall, while on sideslopes and tops the vegetation is a deciduous woodland with trees of 16-18 m height and sclerophyllous leaves. In Cuban mogotes, the slope forest has a 10- to 16-m high open canopy of deciduous trees with barrel-like trunks and abundant columnar cacti, but can quickly grade to a shrubland dominated by terrestrial bromeliads and diverse sclerophyllous shrubs and trees.

Floristics: The following list of species are among those that are diagnostic for this division: Astronium graveolens, Bombacopsis quinata, Bursera simaruba, Bursera fagaroides var. elongata (= Bursera odorata), Bursera instabilis, Bursera morelensis, Calycophyllum candidissimum, Ceiba aesculifolia, Ceiba acuminata, Celtis ehrenbergiana (= Celtis pallida), Coccoloba diversifolia, Conzattia sericea, Enterolobium cyclocarpum, Erythroxylum areolatum, Eugenia axillaris, Exostema caribaeum, Exothea paniculata, Guazuma ulmifolia, Guettarda krugii, Guaiacum sanctum, Guapira obtusata, Gymnanthes lucida, Hypolate trifoliate, Jarilla heterophylla (= Jarilla chocola), Krugiodendron ferreum, Maclura tinctoria, Martinella obovata (= Tabebuia cordata), Metopium toxiferum, Pinus elliottii var. densa, Plumeria obtusa, Sideroxylon foetidissimum, Sideroxylon salicifolium, and Tabebuia ochracea. Characteristic pine species in the Caribbean include Pinus caribaea (var. bahamensis, var. caribaea, and var. hondurensis), and Pinus tropicalis. Diagnostic shrubs include Chrysobalanus icaco, Coccothrinax argentata, Ilex cassine, Sideroxylon salicifolium, Tetrazygia bicolor, and Thrinax morrisii. Similar open pine forests occur in the northern Bahama Archipelago and along the coast in western and eastern Cuba where they are known as pine barrens or pine woodlands and are dominated by Pinus caribaea, Pinus tropicalis, and Pinus cubensis, respectively, accompanied by a similar set of species such as the palms Coccothrinax argentata, Coccothrinax miraguana, Coccothrinax orientalis, among others. Given the harsh conditions of the Antillean dry forests, those relatively few species that thrive under the stress are able to dominate sites (Lugo et al. 2006). Dominant species in Puerto Rico coastal dry forest are Coccoloba krugii, Erithalis fruticosa, Exostema caribaeum, Guettarda krugii, Gymnanthes lucida, Pisonia albida, Savia sessiliflora, and Thouinia striata var. portoricensis. Endemism in these Caribbean forests is very high and represents about 50% of the characteristic plant species.

The division also includes pine-oak dry forests in western Cuba, where the oak species Quercus oleoides also occurs in southern Mexico and Central America. Mesoamerican dry forests occur on soils of variable depth, texture, and alkalinity, and typically have a closed canopy 10-25 m high depending on the type and nutrient contents of the soil. A moderate to high diversity of tropical deciduous tree species dominate the multi-tiered tree canopy. The structure and composition vary along the large distributional range, which includes mountainous topographies that influence moisture availability and soil characteristics. Characteristic genera include Astronium, Bursera, Ceiba, Cassia, Calycophyllum, Cochlospermum, Cordia, Enterolobium, Ficus, Gyrocarpus, Lysiloma, Plumeria, Platymiscium, Pterocarpus, Thouinidium, Zanthoxylum, among others. At the northwestern extreme in Sinaloa, Mexico, these forests include Conzattia sericea, Jarilla heterophylla, Bursera inopinnata, Ceiba acuminata, Martinella obovata, Ipomoea arborescens, Lysiloma watsonii, Choclosperma vitifolium, Pachycereus pecten-aboriginum, Stenocereus thurberi, Mardensia edulis, Senna bicapsularis (= Cassia emarginata), and Tillandsia inflata. Further south from Jalisco through Oaxaca, common species include Achatocarpus oaxacanus, Aphipterygium glaucum, Bombax ellipicum, Bombax palmeri, Bursera fagaroides, Bursera instabilis, Bursera longipes, Bursera morelensis, Bursera fagaroides var. elongata, Ceiba aesculifolia, Coccoloba spp., Pseudosmodingium perniciosum, and Tabebuia palmeri. Dry inter-montane valleys of south-central Mexico (e.g., Balsas dry forest) include Agave pedunculifera, Bursera ariensis, Bursera diversifolia, Bursera hintonii, Ceiba aesculifolia, Cochlospermum vitifolium, Conzattia multiflora, Cordia elaeagnoides, Cyrtocarpa procera, Ficus cotinifolia, Ficus goldmanii, Ficus kellermanni, Ficus petiolaris, Haematoxylon brasiletto, Heliocarpus reticulatus, Lysiloma divaricatum, Pterocarpus orbiculatus, Ruprechtia fusca, Tabebuia impetiginosa, and Vitex pyramidata. Throughout the northern Yucatán Peninsula, most of its distribution is related to limestone outcrops of coral origin, up to hilly terrain and supports a seasonal climate with low annual rainfall. Characteristic species include Beaucarnea pliabilis, Caesalpinia gaumeri, Caesalpinia vesicaria, Diospyros cuneata, Guaiacum sanctum, Hampea trilobata, Lemaireocereus griseus, Lemairocereus aragonii, Lysiloma latisiliquum, Manilkara sapota, Parmentiera aculeata, Plumeria obtusa, Pseudophoenix sp., and Pterocereus gaumeri. Further south along the Pacific Central American coast, extending to the Azuero Peninsula of Panama, characteristic species include Acacia collinsii, Adenocalymma inundatum, Albizia caribea, Allophyllus occidentalis, Andira inermis, Apeiba spp., Ardisia revoluta, Arrabidaea mollissima, Astronium graveolens, Bauhinia glabra, Bombacopsis quinata, Bursera simaruba, Calycophyllum candidissimum, Casearia arguta, Cedrela odorata, Ceiba aesculifolia, Chomelia spinosa, Cochlospermum vitifolium, Combretum farinosum, Cydista diversifolia, Enterolobium cyclocarpum, Exostema mexicanum, Ficus spp., Genipa americana, Guaiacum sanctum, Guarea excelsa, Guazuma ulmifolia, Gyrocarpus americanus, Hemiangium excelsum, Jacquinia pungens, Lonchocarpus phaseolifolius, Lonchocarpus phlebophyllus, Luehea candida, Maclura tinctoria, Platymiscium pleiostachyum, Sabal allenii, Sideroxylon capiri (= Mastichodendron capiri), Simarouba amara, Simarouba glauca, Spondias mombin, Stemmadenia obovata, Sterculia apetala, Swietenia humilis, Swietenia macrophylla, Tabebuia ochracea, Thouinidium decandrum, Trichilia colimana, and Zanthoxylum setulosum.

Naturalized species that may be common or may dominate ruderal dry forests include Vachellia farnesiana (= Acacia farnesiana), Leucaena leucocephala, Melicoccus bijugatus, Parkinsonia aculeata, Prosopis juliflora, and Tamarindus indica. Some of the most widespread exotic trees and shrubs in Puerto Rico and the Virgin Islands are Albizia lebbeck, Leucaena leucocephala, Pinus caribaea, Schinus molle, Spathodea campanulata, Syzygium jambos, Tabebuia rosea, and Terminalia catappa (Brandeis et al. 2009b, Chinea and Helmer 2009).

Dynamics:  Caribbean and Mesoamerican dry forests tend to be exposed to harsh environmental conditions that, depending on their intensity, can cause damage or diebacks, such as seasonal water deficit, nutrient stress, hurricane-force winds and salt spray, and saltwater storm surge. This has influenced the development of structural and physiological mechanisms, making them resilient to disturbance. Among the more outstanding ones are a high resistance to wind (short stature), a high proportion of root biomass, high soil carbon and nutrient accumulation below ground, the ability of most tree species to resprout, and high nutrient use efficiency (Lugo et al. 2006). Fire is not thought to be part of the natural dynamics of these dry forests, but hurricanes often are, which naturally results in considerable heterogeneity in habitat structure and food availability on small spatial scales. This structuring of coastal dry forest by frequent natural disturbance may favor their resilience to anthropogenic disturbance and fragmentation.
Environmental Description:  This division includes tropical forests characterized by a dry season of several months occurring close to the coast or in interior valleys throughout Mexico, Central America, the Caribbean Basin, and south Florida. There, mean annual temperatures in the area of distribution range from 23°C (74°F) in the north to 26°C (77°F) in the Lower Keys. Caribbean dry forests occurring on the mainland in Florida and the Upper Keys are periodically exposed to short-term frost and their flora is composed of a subset of native tropical trees that can withstand rare frost events. Mean annual temperature in the West Indies distribution is around 25°C. Precipitation primarily occurs from May or June to October and ranges from 1650 mm along the Atlantic coast decreasing southward to less than 1000 mm in the Lower Keys (Gillespie 2006). Annual precipitation in the distributional range of this forest in Cuba is less than 1500 mm in the west part of the range and increases towards the east. Precipitation in the distribution range of this forest in Puerto Rico and over most of the islands of Culebra and Vieques ranges from 600 to 1100 mm per year (Brandeis et al. 2006), with two dry seasons, the longer one from December to April and a shorter one from June to August. Throughout Mexico and Central America, these seasonally dry deciduous to semi-deciduous forests are distributed from sea level up to 1400 m elevation in the Pacific basin from Mexico to Panama. Precipitation generally varies from 600 to 1600 mm/year and annual mean temperatures are over 24°C. These forests experience at least one distinct dry season of 4-6 months/year.

Dry forests may often be exposed to highly stressful conditions given the combination of environmental features such as low moisture availability, long dry seasons, decadal cycles of pronounced drought, wind exposure and salt spray in littoral locations. These forests are also periodically exposed to hurricane conditions, with effects that span from flooding with seawater to defoliation, treefall and other structural changes due to strong winds.

Limestone is the dominant substrate in Caribbean dry forests, and in the Yucatán Peninsula of Mexico, with skeletal organic soils with minor mineral components, rarely exceeding 20 cm in depth (Snyder et al. 1990, cited in Gillespie 2006). In the Greater Antilles the distribution of dry forests is indicative of limestone substrates occurring in narrow strips on the northern and southern coastal areas. These also include karst formations with steep slopes and plateaus of towerlike karstic hills up to 300-600 m elevation, with bare karstic rock or more-or-less eroded skeletal soils, or limestone cliffs, and the narrow valleys and gorges in between. Rocky limestone soils have low water-holding capacity and nutritional limitations imposed by their calcareous composition. In flat low-lying limestone archipelagos, such as the Bahamas, the Cayman Islands, Mona and Anegada, dry forests and shrublands dominate. In volcanic, low mountainous islands of the Lesser Antilles, dry forests dominate except for protected sites and ravines where moist forest can grow (Lugo et al. 2006). In Cuba, the dry pine forests are found primarily on acidic soils that have little water-retention capacity and are poor in essential elements. The principal soil types on which they occur are quartziferous sands, pseudo-spodosols in the west and lateritic soils in the east. Only pine trees, which have an ectomycorrhizal symbiosis with fungi, are capable of obtaining in this way a sufficient amount of nutrients to achieve the size of trees. In the Bahamas, pine rockland occurs on relatively flat, moderately to well-drained terrain, from 2-7 m above sea level (Snyder et al. 1990). The oolitic limestone is at or very near the surface, and there is very little soil development. Soils are generally composed of small accumulations of nutrient-poor sand, marl, clayey loam, and organic debris in depressions and crevices in the rock surface. Drainage varies according to the porosity of the limestone substrate, but is generally rapid. Consequently, most sites are wet for only short periods following heavy rains. During the rainy season, however, some sites may be shallowly inundated by slow-flowing surface water for up to 60 days each year (FNAI 2010a).

Dry forests throughout Mexico and Central America occur across a wider diversity of local climate conditions, landforms, and substrates than in the Caribbean. In Tamaulipas, Mexico, these forests occur across lower elevation slopes of the eastern Sierra Madre Oriental, Sierra de San Carlos, Sierra de Tamaulipas, and plateaus of northeastern Mexico. In Sinaloa they are found in canyons and steep slopes with thin, sandy soils. These areas typically experience two wet seasons (winter and mid-summer) and two dry seasons each year. This favors deciduous plant species with well-developed root storage systems, able to rapidly respond to wet seasons. In Nayarit these forests occur at elevations from sea level up through 1,300m, always on slopes. These forests occur on soils of variable depth, texture, and alkalinity. Throughout Guerrero they are found in canyons and steep slopes with soils that are shallow, sandy, nutrient poor, and have low water holding capacity. These areas (extending south through Central America) typically experience one distinct wet season (July-November) and one dry season (December-June) each year. In Guerrero, this vegetation occurs between 0 - 900 m elevation with mean annual precipitation varying between 400-1200 mm, and mean annual temperature around 25oC. These communities transition into thorn scrub at the drier end of its gradient along lower elevation borders, and to semi-evergreen forest further upslope or at the valley bottoms. In the Motagua Valley of Guatemala, and nearby valleys in Honduras, this vegetation usually occurs on deep, rich soils with annual rainfall between 1000 and 1500 mm, but can reach up to 2000 mm and an average temperature of 24oC. With increasing elevation surrounding these valleys, there is an abrupt transition to premontane wet forests. Throughout the remaining Pacific coast of Central America, these forests grow on a variety of soils and topography, with substrates often derived from volcanic activity or from limestones or sandstones. In general, these tend to be more fertile soils than in other parts of the range of this division. Weather conditions that characterize this southerly range are more constant, with an average annual precipitation ranging up to 2000 mm, always with a dry season of 4-6 months, but the exact period varies according to locality.
Geographic Range: This division is found throughout the Caribbean, south Florida, Caribbean coastal South America, Gulf Coast of Mexico and Pacific Coast of Mexico and Central America.

Nations: BS,BZ,CO,CR,CU,DO,GT,HN,HT,JM,KN,MQ,MX,NI,PA,PR,SV,TC,TT,US,VE,VG?,VI,XC,XD

States/Provinces:  FL


Confidence Level: High

Confidence Level Comments: No Data Available
Grank: GNR

Greasons: No Data Available
Type Name Database Code Classification Code
Class 1 Forest & Woodland Class C01 1
Subclass 1.A Tropical Forest & Woodland Subclass S17 1.A
Formation 1.A.1 Tropical Dry Forest & Woodland Formation F003 1.A.1
Division 1.A.1.Ea Caribbean-Mesoamerican Dry Forest & Woodland Division D099 1.A.1.Ea
Macrogroup 1.A.1.Ea.1 Florida Silver Palm - Florida Slash Pine - Caribbean Pine Dry Forest Macrogroup M296 1.A.1.Ea.1
Macrogroup 1.A.1.Ea.2 Gumbo Limbo - Tie-tongue - Stopper species Caribbean Coastal Lowland Dry Forest Macrogroup M134 1.A.1.Ea.2
Macrogroup 1.A.1.Ea.3 Caribbean Dry Limestone Forest Macrogroup M294 1.A.1.Ea.3
Macrogroup 1.A.1.Ea.90 Caribbean Ruderal Dry Forest Macrogroup M514 1.A.1.Ea.90
Concept Lineage: No Data Available

Predecessors: No Data Available

Obsolete Names: No Data Available

Obsolete Parents: No Data Available
Synonomy: No Data Available
Concept Author(s): C. Josse and P. Comer, in Faber-Langendoen et al. (2015)

Author of Description: P. Comer and C. Josse

Acknowledgements: J. Franklin

Version Date: 01-07-16
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