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M134 Bursera simaruba - Coccoloba diversifolia - Eugenia sp. Caribbean Coastal Lowland Dry Forest Macrogroup
Type Concept Sentence: This vegetation is found in flat lowlands and low hills (ca. 300 m elevation) with a pronounced dry season along south Florida and the Antilles. Forests have low stature, high density of small and medium-sized trees, and have single-layer canopies with no emergent trees. The herb layer is poorly developed or completely lacking. Floristic diversity is low compared to more humid forests, and high species dominance is common. Variation in the dry season period, topography, and substrate determine the specific characteristics of the vegetation communities in this macrogroup since all of these have great importance in dry forests as determinants of variation in water availability.
Common (Translated Scientific) Name: Gumbo Limbo - Tie-tongue - Stopper species Caribbean Coastal Lowland Dry Forest Macrogroup
Colloquial Name: Caribbean Coastal Lowland Dry Forest
Hierarchy Level: Macrogroup
Type Concept: This macrogroup encompass tropical and subtropical forests characterized by a dry season of several months that occur in coastal lowlands and low hills, littoral or sublittoral flatlands with rock outcrops and higher terraces facing the sea, on limestone coral shelves, humic carbonate soils, shallow red ferralitic soils, or sandy soils of stabilized, old coastal dunes in south Florida, the Florida Keys, the Greater Antilles, and other Caribbean islands such as those of the Bahamas and Virgin Islands archipelagos. The species composition and structure of these forests vary depending upon the substrate and climate across their distribution. They are broadleaf semi-deciduous to evergreen forests with a canopy between 6-10 m of height. The density of stems tends to be very high. The woody understory is mostly evergreen. The herb layer is poorly developed or completely lacking. In the continental U.S., the Caribbean dry forest occurs in two settings: the hardwood hammock forest in southern Florida, on elevated outcrops of limestone, and the strand forest - a narrow band of hardwood forest and tall shrublands lying just inland of the coastal dune system in south Florida. The latter occur on stabilized, old coastal dunes, often with substantial shell components. In both cases the vegetation is characterized by hardwood species with tropical affinities, with Eugenia axillaris as the most commonly shared species. The Florida hammock forest occurs in three discrete regions, including the Florida Keys, southeastern Big Cypress, and the Miami Rock Ridge. Tropical hardwood species likely to be encountered include Ardisia escallonoides, Bursera simaruba, Coccoloba diversifolia, Eugenia axillaris, and Guapira discolor. The northward ranges of these species are limited by the incidence of frost. These forests tend to have a dense canopy that produces deeper shade, less evaporation, and lower air temperature than surrounding vegetation in these locations. This microclimate, in combination with high water tables, tends to keep humidity levels high. A number of orchid and bromeliad species thrive in such conditions. Unlike much U.S. coastal plain vegetation, fire is a major threat to these hardwood hammock forests.
Dry forests of the Caribbean islands share some features with the Florida dry forests, such as the high density of stems, low stature and relative low floristic diversity, with several shared species among the dominant ones (Bursera simaruba, Coccoloba diversifolia, Eugenia foetida, Gymnanthes lucida). Diagnostic species of this type include Amyris elemifera, Bursera simaruba, Bucida buceras, Coccoloba diversifolia, Coccoloba uvifera, Coccoloba krugii, Eugenia axillaris, Eugenia foetida, Erithalis fruticosa, Exostema caribaeum, Haematoxylum campechianum, Gymnanthes lucida, Guettarda krugii, Guaiacum officinale, Guaiacum sanctum, Jacquinia armillaris, Krugiodendron ferreum, Nectandra coriacea, Pisonia albida, Sabal palmetto, Simarouba glauca, Savia sessiliflora, and Thouinia striata var. portoricensis. These forests also share the limestone substrate which is widespread among the coastal Antilles. Caribbean coastal dry forests in Cuba are slightly taller and have two canopy layers; with the upper layer reaching 12-15 m. This macrogroup also includes lowland semi-deciduous forests on richer substrates in Cuba, Hispaniola, and several smaller Caribbean islands. However, centuries of human occupation and agricultural land use have largely replaced these natural forests described in the literature as more diverse, semi-deciduous, and with the canopy up to 15-20 m height.
Dry forests of the Caribbean islands share some features with the Florida dry forests, such as the high density of stems, low stature and relative low floristic diversity, with several shared species among the dominant ones (Bursera simaruba, Coccoloba diversifolia, Eugenia foetida, Gymnanthes lucida). Diagnostic species of this type include Amyris elemifera, Bursera simaruba, Bucida buceras, Coccoloba diversifolia, Coccoloba uvifera, Coccoloba krugii, Eugenia axillaris, Eugenia foetida, Erithalis fruticosa, Exostema caribaeum, Haematoxylum campechianum, Gymnanthes lucida, Guettarda krugii, Guaiacum officinale, Guaiacum sanctum, Jacquinia armillaris, Krugiodendron ferreum, Nectandra coriacea, Pisonia albida, Sabal palmetto, Simarouba glauca, Savia sessiliflora, and Thouinia striata var. portoricensis. These forests also share the limestone substrate which is widespread among the coastal Antilles. Caribbean coastal dry forests in Cuba are slightly taller and have two canopy layers; with the upper layer reaching 12-15 m. This macrogroup also includes lowland semi-deciduous forests on richer substrates in Cuba, Hispaniola, and several smaller Caribbean islands. However, centuries of human occupation and agricultural land use have largely replaced these natural forests described in the literature as more diverse, semi-deciduous, and with the canopy up to 15-20 m height.
Diagnostic Characteristics: Diagnostic characteristics for this forest macrogroup are low stature with few or no emergent trees, poorly distinguished canopy layers, a high density of stems resulting from the majority of tree species having a tendency to develop several main stems (to coppice), and a dry season severe enough to select for drought-deciduous or evergreen, drought-tolerant trees. Among the species likely to be encountered throughout are Bursera simaruba, Coccoloba diversifolia, and Eugenia axillaris. The tree species of the tropical hammocks of Florida are a subset of the native Caribbean dry forests species that can withstand extremely rare frost events (Gillespie 2006).
Rationale for Nominal Species or Physiognomic Features: No Data Available
Classification Comments: The distinction between this macrogroup and ~Caribbean Dry Limestone Forest Macrogroup (M294)$$ should be clarified, since the latter refers to plant communities that grow on limestone substrates in dry climates, but also in other dry geophysical settings within humid climates. Calcium carbonate-rich soils create physiological and nutrient stress on the plant communities growing on them which results in distinct structure and composition.
Similar NVC Types: No Data Available
note: No Data Available
Physiognomy and Structure: The dry forests of south Florida and the Caribbean have a greater density of individual stems and shorter canopy heights than tropical dry forests in the mainland Neotropics (Gillespie 2006, Lugo et al. 2006). Density varies from 14,000 stems >2.5 cm/ha in Puerto Rico dry forest to 4600 stems/ha in Florida. Canopy height ranges between 5 and 10 m, with a higher mean height in Florida hammock forest stands than in Puerto Rico dry forest, and few trees reaching 18-20 m height. The canopy is seasonally open; however, there are few 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. 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). There is a notable lack of lianas compared with dry forests in the mainland Neotropics (Gillespie 2006). Despite relatively low species diversity, 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). In south Florida, dry tropical forests occurring in sites with higher mean temperature and lower precipitation have greater stand density, while higher precipitation and lower temperatures result in greater canopy heights (Gillespie 2006). Epiphytic orchid and bromeliad species are often found in areas where frosts or anthropogenic disturbances have not occurred in a long time.
Floristics: The vegetation is characterized by hardwood species with tropical affinities. Common species in south Florida, the Bahamas, Cuba and Puerto Rico are Amyris elemifera, Bursera simaruba, Coccoloba diversifolia, Coccoloba uvifera, Eugenia axillaris, Eugenia foetida, Gymnanthes lucida, Krugiodendron ferreum, Nectandra coriacea, Sabal palmetto, and Simarouba glauca. 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. 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). Endemism is very high and represents about 50% of the species. The northern extent of this macrogroup is limited by periodic freezes and cold-tolerance of tropical constituent species, such as Ardisia escallonoides, Eugenia axillaris, Exothea paniculata, Guapira discolor, Metopium toxiferum, Nectandra coriacea, and Piscidia piscipula (Johnson and Muller 1993a).
Dynamics: Along the Florida distribution of this macrogroup, the coastal maritime hammocks are prone to disturbance from hurricanes, which can include extremely high winds, and in coastal areas salt spray, and saltwater storm surge. The rockland habitats on the Florida mainland are mostly inland and not subject to the salt spray and storm surge associated with hurricanes. However, winter freezes sometimes occur and result in damage of the tropical woody plants. In the Florida Keys, freezing temperatures are very unusual.
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). There is a notable lack of lianas compared with dry forests in the mainland Neotropics (Gillespie 2006). Despite relatively low species diversity, 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). Caribbean dry forests have to cope with 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.
Overall, Caribbean coastal dry forests are exposed to harsh environmental conditions that, depending on their intensity, can cause damage or diebacks, such as seasonal water deficit, nutrient stress, strong winds and salt spray, and saltwater storm surge. This has influenced the development of structural and physiological mechanisms to cope, making them very 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 part of the natural dynamics of Caribbean coastal dry forests (though many dry forests are now subject to anthropogenic fires). This is why many examples occur alongside natural firebreaks, such as the leeward side of exposed limestone (Robertson 1955), moats created by limestone solution (Duever et al. 1986), and elevated outcrops above marshes, scrub cypress, or sometimes mangrove swamps (Snyder et al. 1990), or isolated on ridges in pine woodlands.
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). There is a notable lack of lianas compared with dry forests in the mainland Neotropics (Gillespie 2006). Despite relatively low species diversity, 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). Caribbean dry forests have to cope with 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.
Overall, Caribbean coastal dry forests are exposed to harsh environmental conditions that, depending on their intensity, can cause damage or diebacks, such as seasonal water deficit, nutrient stress, strong winds and salt spray, and saltwater storm surge. This has influenced the development of structural and physiological mechanisms to cope, making them very 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 part of the natural dynamics of Caribbean coastal dry forests (though many dry forests are now subject to anthropogenic fires). This is why many examples occur alongside natural firebreaks, such as the leeward side of exposed limestone (Robertson 1955), moats created by limestone solution (Duever et al. 1986), and elevated outcrops above marshes, scrub cypress, or sometimes mangrove swamps (Snyder et al. 1990), or isolated on ridges in pine woodlands.
Environmental Description: Climate: 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 temperatures in the Caribbean dry forest region range from 23°C (74°F) in the north to 26°C (77°F) in the Lower Keys. Precipitation primarily occurs from 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). 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. 2009), with two dry seasons. U.S. Virgin Islands examples may be even drier. Some of the U.S. Virgin Islands examples reach 1200 mm per year.
Soil/Substrate: Limestone is the dominant substrate in Caribbean dry forests of Florida and the Bahamas, with skeletal organic soils with minor mineral components, rarely exceeding 20 cm in depth (Snyder et al. 1990, as cited in Gillespie 2006). In Florida, the coastal maritime hammocks are found on stabilized coastal dunes, often with substantial shell components. 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. Isolated inland, ultramafic soils associated with limestone also support dry forests. 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).
Caribbean dry forests have to cope with 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.
Soil/Substrate: Limestone is the dominant substrate in Caribbean dry forests of Florida and the Bahamas, with skeletal organic soils with minor mineral components, rarely exceeding 20 cm in depth (Snyder et al. 1990, as cited in Gillespie 2006). In Florida, the coastal maritime hammocks are found on stabilized coastal dunes, often with substantial shell components. 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. Isolated inland, ultramafic soils associated with limestone also support dry forests. 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).
Caribbean dry forests have to cope with 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.
Geographic Range: This macrogroup is found in south Florida, the Florida Keys, the Bahamas, Cayman Islands, Cuba, Hispaniola, Jamaica, Leeward Islands, Puerto Rico, Trinidad and Tobago, and Windward Islands.
Nations: BS,CU,DO,HT,JM,MQ,PR,TC,TT,US,VE,VG?,VI,XD
States/Provinces: FL
Plot Analysis Summary:
http://vegbank.org/natureserve/ELEMENT_GLOBAL.2.838402
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.2 Gumbo Limbo - Tie-tongue - Stopper species Caribbean Coastal Lowland Dry Forest Macrogroup | M134 | 1.A.1.Ea.2 |
| Group | 1.A.1.Ea.2.a Caribbean Coastal Dry Evergreen Forest Group | G476 | 1.A.1.Ea.2.a |
| Group | 1.A.1.Ea.2.b Gumbo Limbo - Tie-tongue - White Stopper Forest Group | G765 | 1.A.1.Ea.2.b |
Concept Lineage: No Data Available
Predecessors: No Data Available
Obsolete Names: No Data Available
Obsolete Parents: No Data Available
Synonomy: = Rockland Forest (Ross et al. 1992)
= Tropical Hammock (Snyder et al. 1990)
= Tropical Hammock (Snyder et al. 1990)
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