Distribution of Vegetation Types

Abstract

This chapter presents an overview of the vegetation types of the Yucatán Peninsula. Mean annual precipitation, terrain and soil characteristics explain the large-scale distribution of forest types in the Yucatán Peninsula. Tropical (high) forest is found in areas with >800 mm of mean annual precipitation, while dry (low) tropical forest is distributed in areas with <800 mm of mean annual precipitation. Tropical forest has canopy heights of more than 30 m and mostly presents three well-defined vegetative strata. The distribution of tropical forest is mainly in the central, eastern and southern parts of the Yucatán Peninsula. Dry tropical forest has canopy heights up to 20 m, and is widely distributed in the Yucatán Peninsula in different successional stages. Other woody vegetation types include mangroves, petenes and pine savannas. Open vegetation types include coastal dunes, marsh, and savanna vegetation. Disturbed tropical and dry tropical forest of all successional stages covers more than 7.4 million ha at present, and requires detailed management plans to maintain future ecosystems benefits.

Full text

Chapter 3
Distribution of Vegetation Types
Gerald Alexander Islebe, Odil
on Sa
´nchez-Sa
´nchez,
Mirna Valde
´z-Herna
´ndez, and Holger Weissenberger
Abstract This chapter presents an overview of the vegetation types of the Yucata
´n
Peninsula. Mean annual precipitation, terrain and soil characteristics explain the
large-scale distribution of forest types in the Yucata
´n Peninsula. Tropical (high)
forest is found in areas with >800 mm of mean annual precipitation, while dry
(low) tropical forest is distributed in areas with <800 mm of mean annual precipi-
tation. Tropical forest has canopy heights of more than 30 m and mostly presents
three well-defined vegetative strata. The distribution of tropical forest is mainly in
the central, eastern and southern parts of the Yucata
´n Peninsula. Dry tropical forest
has canopy heights up to 20 m, and is widely distributed in the Yucata
´n Peninsula in
different successional stages. Other woody vegetation types include mangroves,
petenes and pine savannas. Open vegetation types include coastal dunes, marsh, and
savanna vegetation. Disturbed tropical and dry tropical forest of all successional
stages covers more than 7.4 million ha at present, and requires detailed manage-
ment plans to maintain future ecosystems benefits.
Keywords Tropical forest • Distribution • Precipitation • Succession •
Mangroves • Coastal dunes • Marsh vegetation
3.1 Introduction
Although the Yucata
´n Peninsula is characterized as an extensive limestone plateau
with no significant altitudinal variation, a relatively large number of vegetation types
can be identified. Vegetation types are mainly distributed along a north-to-south
G.A. Islebe (*) • M. Valde
´z-Herna
´ndez • H. Weissenberger
El Colegio de la Frontera Sur Unidad Chetumal, Avenida Centenario km 5.5, Apartado Postal
424, CP 77014 Chetumal, Quintana Roo, Mexico
e-mail: gislebe@ecosur.mx;mavaldez@ecosur.mx;hweissenberger@ecosur.mx
O. Sa
´nchez-Sa
´nchez
Universidad Veracruzana, Centro de Investigaciones Tropicales de la Universidad
Veracruzana, Ex Hacienda Lucas Martı
´n, Privada de Araucarias s/n, Col. Periodistas, 91019
Xalapa, Veracruz, Mexico
e-mail: sasodil@yahoo.com.mx
©Springer International Publishing Switzerland 2015
G.A. Islebe et al. (eds.), Biodiversity and Conservation of the Yucat
an Peninsula,
DOI 10.1007/978-3-319-06529-8_3
39

gradient of precipitation and according to different soil types, as well as precipitation
variability in an east to west direction (Miranda 1958; Wright 1967). Tropical forest
vegetation can be classified into two major groups with stand-specific associations,
namely high and low forest. Other forest types, like mangrove forest, are found
along coastal areas of the peninsula. Aquatic and subaquatic vegetation is present in
open seasonally flooded areas locally known as sabanas, swamps, lakes and other
water bodies.
The climate and hence the vegetation of the Yucata
´n Peninsula is determined by
a series of factors: the lack of major orographic variation, the Tropic of Cancer,
the influence of the Bermuda High, the presence of jet streams, tropical cyclones
during the rainy season, cold fronts during the winter months, and the presence of a
warm ocean current in the Yucata
´n Channel (Orellana et al. 2003). The driest region
is located in the northwestern Yucata
´n Peninsula, in the Sisal-Progreso region,
where mean annual precipitation is around 500 mm. The isohyet of 1000 mm runs
east to west from northern Campeche, Yucata
´n to northern Quintana Roo. Toward
the southern part of the Gulf of Mexico, precipitation increases rapidly up to
2000 mm; to the south of Quintana Roo, isohyets of around 1400 mm are found.
Most of the precipitation falls between May and November, while the period
between December and April is considered dry. Cold fronts between December
and February are locally known as nortes, and provide occasional winter
precipitation.
Tropical forest is found as the potential natural vegetation in areas
with >800 mm of annual precipitation, mainly in the central, eastern and southern
Yucata
´n Peninsula, in the states of Campeche and Quintana Roo. High forest as
a major vegetation type presents floristic and structural variations and covers
around 70 % of the peninsula before significant human-induced landscape trans-
formation, which began in the twentieth century (Rogan et al. 2011; Schmook
et al. 2011).
Tropical dry forest is generally found in areas with generally less than 800 mm
average precipitation, on a variety of soil types, including Gleysols to Vertisols, and
as a potential natural vegetation type, covers large parts of Quintana Roo, Campe-
che and Yucata
´n. A unique vegetation type is low flooded forest, covering
low-lying areas in Quintana Roo and Campeche. Species of this vegetation type
withstand several months of flooding, as well as dry conditions. Petenes, also a
unique vegetation type, are forest islands with fresh water inlets surrounded by
marsh vegetation.
Recent human impact has caused profound changes in vegetation (Rico-Gray
and Garcı
´a-Franco 1991) due to over-exploitation of timber species and changes in
land use (Rueda 2010; Snook and Negreros 2004). Many plant species have specific
ethnobotanical uses and values, ranging from medicinal to religious purposes
(Barrera 1962; La Torre-Cuadros and Islebe 2003; Andersen et al. 2005). The
Yucata
´n Peninsula is recognized to have an exceptional degree and detail of
traditional ecological knowledge and utilization of local plants.
40 G.A. Islebe et al.

3.2 Forest Types and Distribution
The first systematic descriptions of the vegetation of the Yucata
´n Peninsula are
provided by Miranda (1958), Miranda and Herna
´ndez-X (1964) and Barrera (1962).
Earlier work by Lundell (1934,1937) compiled botanical and ecological aspects of
the vegetation of northern Guatemala and the Yucata
´n Peninsula. The work of
Lundell was impressive, as he also included ethnobotanical and economic factors in
his reports, mainly from observations and data collection in Campeche, Yucata
´n
and the Pete
´n province of northern Guatemala. Lundell’s botanical collections can
be seen in Chicago’s Field Museum of Natural History (http://collections.mnh.si.
edu/search/botany/). Lundell (1937) divided the vegetation in two categories,
upland and lowland, which correspond to specific soil characteristics. The upland
type included broad-leaved forest, also known as “high forest” or (seasonally)
evergreen tropical forest, the soils of which featured a distinct organic layer.
Lowland vegetation types were characterized by Lundell as forest with clayey
soils originating in erosion from the uplands. This vegetation type is locally
known as akalche´.
In the classic work “Vegetaci
on de Me
´xico”, Rzedowski (1978) described
species composition and physiognomy of the major vegetation types of the Yucata
´n
Peninsula. Other regional vegetation studies are from Sa
´nchez-Sa
´nchez and Islebe
(2002), analyzing plant communities along precipitation gradients, and Barber and
colleagues (1999), presenting phytosociological units. Ibarra-Manrı
´quez and
co-authors (1995) analyzed the phytogeographical relationships of the trees of the
Yucata
´n Peninsula and concluded that the strongest ecological and botanical
affinities exist with other Mesoamerican locations, while affinities with the Carib-
bean region are much weaker.
Distribution of vegetation types are presented in two maps (Figs. 3.1 and 3.2)
based on inventories carried out in 2009, 2010 and 2011 (Inegi 2013). Vegetation
types are separated on the two maps into forest (zonal) vegetation, and vegetation
related to water bodies (azonal). Area covered and geographical distribution is
given in Table 3.1. Disturbed dry tropical forest is the largest category with more
than 3.9 million ha, followed by disturbed tropical forest with over 3.5 million
ha. Following the data of Inegi, tropical and dry tropical forests together occupy
less than 240,000 ha. However, it should be noted that disturbed tropical and dry
tropical forests include all successional stages, from 5 to 30 or more years of age.
Vegetation related to water bodies with fluctuating water tables covers more than
2,474,244 ha in the Yucata
´n Peninsula. Most of these areas are covered by Typha-
and Cyperaceae-dominated swamps and disturbed low flooded forest. Mangrove
forests cover more than 400,000 ha along the coasts of the peninsula. However
increased deforestation rates of nearly 1 % per year (Hirales-Cota et al. 2010;
Sa
´nchez-Sa
´nchez et al. 2009), specifically along the Caribbean coast, are threaten-
ing this ecosystem.
3 Distribution of Vegetation Types 41

3.2.1 Tropical Forest
Tropical forest or high forest (Selva alta and selva mediana subperennifolia
in the Spanish literature) is widely distributed in the Yucata
´n Peninsula
Fig. 3.1 Distribution of main vegetation types
Fig. 3.2 Distribution of vegetation related to water bodies
42 G.A. Islebe et al.

(Sa
´nchez-Sa
´nchez 2000). This vegetation type is characterized by 25–50 % of the
trees composing the high forest losing their leaves during the dry season. Mean
annual precipitation values are above 1200 mm. Mean annual temperature in these
forests is above 20 C, while mean minimum annual temperatures are 11 C. Trees
have canopy heights of more 30 m and three well-defined strata can generally be
observed (Martı
´nez and Galindo-Leal 2002).
The dominant tree is Manilkara zapota (Sapotaceae), with a wide distribution in
southern Mexico, extending into northern Belize and Guatemala. Tropical forest is
common in southern-central Campeche and most parts of Quintana Roo. Character-
istic and commercially valuable canopy species include Swietenia macrophylla
(Meliaceae), Swartzia cubensis (Leguminosae), Pimenta dioica (Myrtaceae),
Aspidosperma megalocarpon (Apocynaceae), Caesalpinia gaumeri (Leguminosae),
among many others (Snook and Negreros-Castillo 2004). An upper shrub layer
consists mainly of the species Ficus spp. (Moraceae), Bursera simaruba
(Burseraceae), Sickingia salvadorensis (Rubiaceae), Chlorophora tinctoria
(Moraceae), Enterolobium cyclocarpum (Leguminosae), Sapindus saponaria
(Sapindaceae) and Acrocomia mexicana (Arecaceae). High forest consists of differ-
ent plant communities, in which Manilkara zapota-Coccothrinax readii is the
dominant (Sa
´nchez-Sa
´nchez and Islebe 2002).
Another widely distributed plant community of the high forest is Vitex gaumeri-
Caesalpinia gaumeri. This community has as characteristic species Acacia gaumeri
Table 3.1 Main vegetation types, land use and areas covered 2011
Main vegetation type and land use Cover (ha)
a
Distribution
Agriculture 724,880 PY
Human settlement 149,358 PY
Pastureland 1,561,832 N, C
Tropical forest 1,356,095 E, C, S
Disturbed tropical forest 3,638,819 E, C, S
Dry tropical forest 92,221 W, C, N, S
Disturbed dry tropical forest 3,907,388 W, C, N, S
Water bodies 286,045
Mangrove 400,124 E, N, W
Palm vegetation 6831 E
Typha-swamp vegetation 516,159 E, N, W
Low flooded forest 513,673 E, N, W
Disturbed low flooded forest 656,360 E, N, W
Coastal dune vegetation 8525 E, N, W
Halophytic vegetation 14,392 E, W
Tropical evergreen seasonal alluvial forest 57,176 E
Disturbed mangrove 15,673 E, N, W
Disturbed tropical evergreen seasonal alluvial forest 5339 E
Undefined vegetation types 500,000 PY
PY Yucata
´n Peninsula, NNorth, CCentral Yucata
´n Peninsula, EEast, SSouth, WWest
a
Cover of all data is based on Inegi (2013)
3 Distribution of Vegetation Types 43

(Leguminosae), Byrsonima bucidaefolia (Malphigiaceae), Alseis yucatanensis
(Rubiaceae), Spondias mombin (Anacardiaceae), Diospyros verae-crucis
(Ebenaceae) and Caesalpinia gaumeri (Leguminosae). An additional characteristic
feature of this plant community are the stone slabs, which can cover up to 45 % of
the soil surface (Sa
´nchez-Sa
´nchez and Islebe 2002). In southern Quintana Roo,
where the wettest conditions are found, the characteristic community is Trichilia
glabra-Brosimum alicastrum-Attalea cohune. This community is a dense forest
with the following characteristic subdominant species: Chrysophyllum mexicanum
(Sapotaceae), Piper sempervirens (Piperaceae), Chlorophora tinctorea (Moraceae),
and Swartzia cubensis (Leguminosea).
The Hampea trilobata-Metopium brownei-Bursera simaruba community is
widely distributed in the central Yucata
´n Peninsula and includes many character-
istic taxa of secondary forests (Sa
´nchez-Sa
´nchez and Islebe 2002) like Nectandra
coriacea (Lauraceae), Sabal yapa (Arecaceae), Bauhinia divaricata
(Leguminosea), Lysiloma latisiliquum (Leguminosae) and Bauhinia jenningsii
(Leguminosae).
Soils of the high forest are generally shallow, but rich in organic material and of
Lithosol-Rendzina or even Luvisol and Vertisol types (Flores 1977, FAO 1988;
Wright 1967), depending on the region. In the southern Yucata
´n Peninsula, soils are
mostly Vertisols, while in the central Yucata
´n Peninsula soils are of the Lithosol-
Rendzina type.
Climbers are common in these forests, and characteristic species are Paullina sp.
(Sapindaceae) and Cardiospermum corindum (Sapindaceae). At local terrain
depressions in high forests, pure stands of Attalea cohune (Arecaceae) can be
found (Quero 1992). These local vegetative communities consist of one distinctive
upper layer 20 m in height, and an understory herbaceous layer, typically with 90 %
cover. At similar conditions of deficient drainage, pure stands of Sabal
mauritiiformis (Arecaceae) can be found in southern Quintana Roo. These palm
communities have total heights of nearly 25 m. High forest has nearly disappeared
in the Yucata
´n Peninsula due to excessive timber extraction, cattle farming and
development of rice plantations in southern Quintana Roo over the last 50 years
(Sa
´nchez-Sa
´nchez et al. 2007; Turner et al. 2001). Old growth forest plots have
estimated values of 103.5 4.4 Mg ha
1
AGB (above ground biomass) (Urquiza-
Haas et al. 2007). However, higher values of 190 Mg ha
1
AGB are additionally
reported from the same authors (Urquiza-Haas et al. 2007). Basal area estimates of
tropical forest are highly variable: values between 11.9 and 45.0 m
2
ha
1
are
reported (Urquiza-Haas et al. 2007; Dickinson et al. 2000; Ceccon et al. 2002;
Gonza
´lez-Iturbe et al. 2002; Lawrence and Foster 2002; La Torre-Cuadros and
Islebe 2003; White and Hood 2004). Most likely, differences are due to the sampled
vegetation type and numerical methods applied.
Selva mediana subcaducifolia (semi-deciduous medium dry forest) can be found
in the states of Yucata
´n, Campeche and in some parts of northern Quintana Roo,
although much of its original vegetation has been removed, mostly for logging,
cattle and agriculture (Zamora-Crescencio et al. 2008). Canopy heights are usually
from 20 up to 35 m. Fifty to 75 % of all trees lose their leaves during the dry season.
44 G.A. Islebe et al.

Soils are shallow with rocky outcrops with a very thin organic upper layer, mostly
less than 7 cm (Sa
´nchez-Sa
´nchez and Islebe 2002). These soils are classified as
Lithosols and Luvisols (Zamora-Crescencio et al. 2008). Characteristic species
include Brosimum alicastrum (Moraceae), Vitex gaumeri (Verbenaceae),
Byrsonima (Malphigiaceae), Lysiloma latisiliquum, among others (Dzib-Castillo
et al. 2014; Zamora-Crescencio 2003). Epiphytes can be found in these forests.
3.2.2 Dry Forest
Low seasonally deciduous forest, or selva baja caducifolia in Spanish, has canopy
heights of less than 15 m, and occurs in areas with annual precipitation varying
between 600 and 800 mm or rarely more, located in the states of Yucata
´n,
Campeche and Quintana Roo. Nearly 100 % of tree species drop their leaves during
the dry season. Characteristic species are Lysiloma bahamensis (Leguminosae),
Baeucarnea pliabilis (Nolinaceae), Gymnopodium floribundum (Polygonaceae),
Cassia alata (Leguminosae), Acacia milleriana (Leguminosae), Mimosa
bahamensis (Leguminosae), Diospyros anisandra (Ebenaceae), Pseudophoenix
sargentii (Arecaceae) and Piscidia piscipula (Leguminosae) (Valdez-Herna
´ndez
et al. 2014). Those species can also grow directly on eroded limestone. On a
community level Sebastiania adenophora-Plumeria obtusa var.cericifolia-Agave
angustifolia was identified as dominant in the eastern Yucata
´n Peninsula (Sa
´nchez-
Sa
´nchez and Islebe 2002).
Seasonal flooded low forest (Selva baja inundable) occurs in areas that are
geographically similar to those of high forest (Miranda and Herna
´ndez 1964), but
is related to sabana areas. These forests are found in central and southern Quintana
Roo, as well as northern Campeche and Yucata
´n. Soils with poor drainage are
characteristic of these dry forests, and soils can withstand high water levels during
long periods. The areas of low drainage are locally known as akalche´s, which are
periodically flooded during the rainy season. Several tree species can be found, the
most conspicuous being Byrsonima crassifolia (Malphigiaceae), Chrysobalanus
icaco (Sapotaceae), Curatella americana (Dilleniaceae), Crescentia cujete
(Bignoniaceae) and Hyperbaena winzerlingii (Menispermaceae) (Corte
´s-Castela
´n
and Islebe 2002;Dı
´az-Gallegos et al. 2002). Canopy heights rarely surpass 10 m.
Some of those low forests can form pure stands, as in the case of Haematoxylon
campechianum (Fabaceae) (common name tasistal), Bucidas buceras
(Combretaceae) (common name pukte) and Metopium brownei (Anacardiaceae)
(Miranda and Herna
´ndez 1964). There is no distinctive herbaceous layer present,
mainly due to the prevalence of seasonal flooding. If undergrowth is present,
species belonging to the Poaceae and Cyperaceae families can be found, like
Eleocharis. Epiphytes of Orchidaceae and Piperaceae are quite common, such as
Encyclia alata, Peperomia spp., and climbers belonging to Dahlbergia
(Leguminosae). Deforestation rates for low forest have been estimated around
1.2 % per year in Calakmul for the last 30 years (Schmook et al. 2011).
3 Distribution of Vegetation Types 45

In the northwestern part of the peninsula seasonally dry tropical forest (low
deciduous forest) with columnar cacti can be found (Miranda and Herna
´ndez 1964).
It is restricted to areas of 600–800 mm of annual precipitation, with no precipitation
occurring during a 7- to 8-month period. The presence of rocky areas is conspi-
cuous, and the soils are described as Lithosols (Thien et al. 1982). Characteristic
endemic species of this vegetation type are: Mammillaria gaumeri,Beaucarnea
pliabilis,Guaiacum sanctum,Pilosocereus gaumeri,Nopalea gaumeri,Nopalea
inaperta and Pterocereus gaumeri.
3.2.3 Mangrove Vegetation
Mangroves can be found along the entire coast of the Yucata
´n Peninsula. Four
mangrove species are distributed within different types of vegetative association:
Rhizophoramangle (Rhizophoraceae), Conocarpus erectus (Combretaceae),
Avicennia germinans (Acanthaceae), and Laguncularia racemosa (Combretaceae).
R. mangle forests form nearly pure stands along the coast, while A. germinans and
L. racemosa are found in mixed stands depending on the degree of soil salinity
(Sa
´nchez-Sa
´nchez et al. 1991). The same authors distinguish four different man-
grove communities. C. erectus is distributed on higher grounds with Gleysols, and
can withstand variation in water table depths. Canopy heights of mangrove forests
do not surpass 6–12 m, and consist of one predominant tree layer. The edges of
mangrove areas are mostly dominated by Acrostichum danaeifolium (Pteridophyta),
Bravaisia tubiflora (Acanthaceae), Cladium jamaicensis and other Cyperaceae
(Islebe and Sa
´nchez 2002; Torrescano-Valle and Islebe 2012). In southern Quintana
Roo, dwarf Rhizophora mangle stands can be found with heights of only up to 1 m
(Valde
´z-Hernandez and Islebe 2011). Soils of mangrove communities are mostly
sandy with clayey material, but can develop deep organic horizons.
3.2.4 Peten Vegetation
Peten vegetation is unique to the Yucata
´n Peninsula. It takes the form of closed
woody vegetation islands consisting of a mosaic of mangroves and tropical forest
species surrounded by salt marshes and mangroves (Dura
´n1987). Mangrove
species can occasionally be found mixed with common tropical forest tree species.
The most prominent and largest petenes are those of the biosphere reserve Los
Petenes in Campeche, but these formations are also present on a smaller scale in the
eastern part of Quintana Roo (Sian Ka’an Biosphere). Canopy heights may reach
12 m. Soils are rich in organic material, deep and slightly saline. Characteristic
species are Conocarpus erectus (Combretaceae), Metopium brownei
(Anacardiaceae), Thrinax radiata (Arecaceae), Bucida buceras (Combretaceae),
H. campechianum (Leguminosae), among others.
46 G.A. Islebe et al.

3.3 Open Vegetation Types
Open vegetation types, including sabana, marsh and coastal dune vegetation are
well defined and widely distributed in the Yucata
´n Peninsula. The coastal dune type
is found along the shoreline of the Yucata
´n Peninsula (Espejel 1984,1986,1987;
Moreno-Casasola and Espejel 1986; Flores and Espejel 1986; Torres et al 2010),
and coastal plant communities are distributed along distinctive ecological gradi-
ents, such as salinity (Islebe, unpublished). Espejel (1987) identified 237 species for
coastal dunes, and cosmopolitan families like Poaceae, Asteraceae and
Leguminosae are the most common. Low non-woody plants (<30 cm high) like
Ambrosia hispida (Asteraceae), Canavalia rosea (Leguminosae), Distichlis spicata
(Poaceae), Ipomoea pes-caprae (Convolvulaceae), Suriana maritima (Surianaceae)
and Ernodea littoralis (Rubiaceae), among other species, are found close to the sea.
A transition occurs to higher elevation plant communities with woody taxa like
Coccoloba uviferae (Polygonaceae), Pouteria campechana (Sapotaceae), Cordia
sebestena (Boraginaceae) and Chrysobalanus icaco (Chrysobalanaceae). Woody
taxa reach up to 6 m high, and sometimes a low shrub layer is present (Sa
´nchez-
Sa
´nchez et al. 1991). Plant species from coastal dunes show typical morphological
adaptations such as thicker leaves and glandules (like S. maritima and E. littoralis).
Altitudinal differences between the highest and lowest parts of the dune along
the coasts of Quintana Roo averages 5 m. Near-shore vegetation includes creeping
species like I. pres-caprae, while dunes with woody taxa show the development
of early shallow soil formation with thin organic layers. Coastal dunes are one of
the most threatened vegetation types of the Yucata
´n Peninsula, with less than
8500 ha remaining, as coastal dunes are used to build hotels and tourism infra-
structure (Lapointe 2011). Coastal dune vegetation is mostly delimited by the
presence of Thrinax radiata palm vegetation, or by swamp like vegetation with
abundant Cyperaceae and a transition to tropical forest types. Marsh vegetation is
influenced by a high level of carbonate dissolved in water and soil, and is relatively
species-poor relative to its biogeographical position. Typha dominguensis, Cyperus
jamaicensis, Scirpus and Carex spp. are the main taxa found in marsh environ-
ments and are locally named tulares or popales. Marsh vegetation is mainly
distributed along the eastern and western margins of the peninsula, though marshes
cover large areas of the northern Yum Balam reserve. Their floristic composition is
determined by nutrient availability, phosphate limitation, and the influence of
salinity. T. dominguensis dominates in nutrient-rich conditions (Rejmankova
et al. 1996).
In southern Quintana Roo and reaching further into Belize and parts of Guate-
mala, relic Pinus caribaea forest can be found (Macario-Mendoza et al. 1998). It is
an open savanna-like vegetation type on sandy soils, with taxa from the Poaceae
dominating the herbaceous layer. Following Chavelas (1981) it is the only pine
species occurring at sea level in Mesoamerica, with an original distribution running
from the Yucata
´n Peninsula through Belize to Nicaragua, along coastal areas.
3 Distribution of Vegetation Types 47

3.4 Human and Natural Impact
Human impact on the vegetation of the region has occurred over the last four millennia
(Rico-Gray and Garcı
´a-Franco 1991;Islebeetal.1996). The earliest corn cultivation
in the Yucata
´n Peninsula and its subsequent impact on vegetation is recorded from
2000 BC on (Arag
on-Moreno et al. 2012; Carrillo-Bastos et al. 2010;Torrescano-
Valle and Islebe 2015), although it could date back as much as 5000 years, as evidence
from Northern Belize suggests. The use of agroforestry is part of ancient Mayan
culturalknowledge and is well established among present-day farmers (Barrera 1962).
The sustainable use of natural resources has been documented in many ethnobotanical
studies (Barrera 1962;Andersenetal.2005). Forest sustainability is incorporated into
agroforestry practices like shifting cultivation, depending on the crops and areas
cultivated, as well as cultivation intensity (Schmook et al. 2011; Valdez-Herna
´ndez
et al. 2014). Fire is an important factor, and has been used since early Maya culture to
clear land (Islebe et al. 1996). For the last 4000 years, and until the present day it is,
intentionally or unintentionally, used to modify the landscape at different scales
(L
opez-Portillo et al. 1990). According to soil and paleoecological data from central
Peten in northern Guatemala, full recovery of tropical forest vegetation from fire can
take up to 80 years (Mu
¨ller et al. 2010). However, using physiognomic criteria and
plant species composition data, secondary vegetation cannot be distinguished from
primary vegetation after 30 years (Miranda 1958;Sa
´nchez-Sa
´nchez and Islebe 2002).
Traditional Mayan nomenclature makes no linguistic distinction between secondary
vegetation of more than 30 years and primary forest (Barrera et al. 1976), but the first
seral stages are well defined in traditional ecological knowledge (Sa
´nchez-Sa
´nchez
et al. 2007). Conversely, Lawrence and Foster (2002) report that biomass from
25 year old secondary forests was only 40 % that of mature forests.
Hurricanes as natural hazards are a major determinant of forest structure in the
Yucata
´n Peninsula (Tanner et al. 1991;Sa
´nchez-Sa
´nchez and Islebe 1999; Ramı
´rez-
Barajas et al. 2012; Rogan et al. 2011; Vandecar et al. 2011). The coastal and
immediate interior vegetation of Quintana Roo is especially vulnerable to hurricanes
as natural disturbances, since coral reefs and mangroves are severely affected and
reduced in geographical extent. Hurricane damage to vegetation includes tree
snapping (complete or partial, by branches), uprooting, defoliation, tree mortality,
strong flooding and eventually, weeks after the event, intensive fires, due to the high
quantities of accumulated woody and foliar debris on the ground (L
opez-Portillo
et al. 1990; Rodrı
´guez-Trejo et al. 2011). During the last century, more than
100 hurricanes have hit the coast of Quintana Roo and the Yucata
´n Peninsula,
damaging mangroves and other forest types (Islebe et al. 2009). The hurricanes
causing the most severe damage were Gilbert in 1989 (Sa
´nchez-Sa
´nchez and Islebe
1999), Isidor in 2003, Emily in 2005 and Hurricane Dean (Islebe et al. 2009;
Ramı
´rez-Barajas et al. 2012), which damaged more than 2 million ha of forests in
2007 (CONAFOR, Ramı
´rez-Barajas et al. 2012). For a high forest in northern
Quintana Roo, Sa
´nchez-Sa
´nchez and Islebe (1999) estimated 4.5 t/ha of fallen
biomass during Hurricane Gilbert. For more details, please see Chaps. 6and 7.
48 G.A. Islebe et al.

3.5 Floristic Diversity
Floristic lists of the Yucata
´n Peninsula include the works of Standley (1930),
Cabrera and Sousa (1983) for Quintana Roo, Sosa and colleagues (1985), Duran
and co-authors (1998), Martı
´nez and Galindo-Leal (2002), Gutierrez-Baez (2003)
for Campeche, and Carnevali and co-authors (2010). The latter authors recognize
some 3000 taxa. Some checklists on a local or regional scale are available, e.g., for
the El Eden reserve in northern Quintana Roo, see Schultz (2005) and for Cozumel,
Te
´llez and Cabrera (1987). Endemic species of the Yucata
´n Peninsula are listed in
Dura
´n and colleagues (1998), which follows Ibarra-Manrı
´quez and co-authors
(1995) in summing up 54 species, most of them with Caribbean phytogeographical
affinities.
More than 500 tree species have been identified for the Yucata
´n Peninsula
(based on the CIQRO-Herbarium data base, 2015). The most species-rich woody
families are Leguminosae, Euphorbiaceae and Rubiaceae. Including all plant life
forms, Fabaceae, Poaceae, Asteraceae and Orchidaceae (Carnevali et al. 2010) are
the most represented families. The genera with most species are Ipomoea, Croton,
Euphorbia and Cyperus, following Carnevali and colleagues (2010).
3.6 Outlook for Future Vegetation Studies
Given the threat to all vegetation types, several actions should be taken to preserve
large connected forest tracts, specifically in the core conservation areas of
Calakmul, central Quintana Roo, Campeche and the Sian Ka’an Biosphere reserve.
Conservation efforts are being made to guarantee genetic connectivity and habitat
range for biodiversity, such as the biological corridor of Sian Ka’an-Calakmul. The
effectiveness of these efforts must be analyzed in future years, as the areas
containing this corridor are also used for the construction of roads and rural
infrastructure. Floristic and structural parameters should be monitored; forests
should be locally recognized as ecosystem service providers to boost local income;
and plant demographic studies should be undertaken to establish scientifically
based rules for long term sustainable timber extraction (Valdez-Herna
´ndez
et al. 2014). Scientific knowledge of natural succession should be included in
management plans to avoid excessive unnecessary exploitation of major timber
species with specific basal area parameters. The last 30 years of history has shown
that the ejido (communally owned land) system has worked well (Porter-Bolland
et al. 2013), though not perfectly, to guarantee the conservation of major forest
tracts on a large regional scale in the central and southern Yucata
´n Peninsula.
Although most of the remaining forests are secondary, at different successional
stages, they cover more than 3.5 million ha, and still harbor a high and valuable
degree of biodiversity (Ramı
´rez-Barajas et al. 2012) and supply rural communities
with income and use-value (Rico-Gray and Garcı
´a-Franco 1991). The following
3 Distribution of Vegetation Types 49

research should be encouraged in the future: understanding the succession of the
different vegetation types; improvement and application of restoration ecology;
accurate and precise assessment of the carbon stocks of distinct forest types; and the
eco-physiology of characteristic tree species of Yucata
´n forests. In the case of
mangrove forests, some species are currently protected by national laws. Continu-
ous development of the tourism industry along coastal areas makes effective
conservation almost impossible, however, as ecosystem functioning is altered
with the construction of roads, drainage and other infrastructure.
Acknowledgments We are grateful to the Consejo Nacional de Ciencia y Tecnologı
´a (Conacyt)
for funding our projects and to El Colegio de la Frontera Sur for access to all facilities provided
over the years.
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