tmpFEA4 TMP

The Animal Forest and Its Socio-ecological
Connections to Land and Coastal

Ecosystems
Jess Ernesto Arias-Gonzlez, Andrea Rivera-Sosa, Jaime Zaldvar-

Rae, Christian Alva-Basurto, and Camilo Corts-Useche
Contents
1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.1 Connectivity in Tropical Seascapes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.2 Global Climate Change Impacts on Animal Forests . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2 Linkages and Complexities of Social and Ecological Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
2.1 Transformations of the Yucatan Peninsula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
3 Human-Derived Fluxes and Animal Forest Habitat Change . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
3.1 Chemical Fluxes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
3.2 Biological Fluxes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
3.3 Physical Fluxes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
4 Management and Conservation of the Animal Forest and Tropical Seascapes . . . . . . . . . . . . . 20
4.1 Social Inclusion in the Management of MPAs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
4.2 Current and Future Management Tools . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
4.3 Governance to Improve the Management of Seascape Connectivity . . . . . . . . . . . . . . . . . 24
5 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
6 Cross-References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
J.E. Arias-Gonzlez (*) A. Rivera-Sosa C. Corts-Useche

Laboratorio de Ecologa de Ecosistemas de Arrecifes Coralinos, Departamento de Recursos del
Mar, Centro de Investigacin y Estudios Avanzados del I.P.N-Unidad Mrida, Mrida, Yucatn,
Mexico
e-mail: earias@cinvestav.mx; andrea.rivera@cinvestav.mx; camilo.cortes@cinvestav.mx
J. Zaldvar-Rae
Coordinacin de Investigacin, Vicerrectora Acadmica, Universidad Anhuac Mayab, Mrida,
Yucatn, Mexico
e-mail: jaime.zaldivar@anahuac.mx
C. Alva-Basurto
Parque Nacional Costa Occidental de Isla Mujeres, Punta Cancn y Punta Nizuc. Comisin
Nacional de reas Naturales Protegidas, Cancn, Quintana Roo, Mexico
e-mail: chr.alva@yahoo.com.mx
# Springer International Publishing Switzerland 2016 1

S. Rossi (ed.), Marine Animal Forests,
DOI 10.1007/978-3-319-17001-5_33-1
2 J.E. Arias-Gonzlez et al.
Abstract
Tropical landscape and seascape systems are intimately linked by complex
ecological relationships that provide environmental services to human societies
located in coastal areas and beyond. Paradoxically, nonsustainable activities from
these human societies are threatening the functions and benets of these systems.
Anthropogenic processes that damage tropical seascapes are rapidly increasing as
a result of coastal and tourism development, increasing human population,
unsustainable economic growth, and extensive transformations of natural land-
scapes. In addition to this overwhelming trend, tropical coastal seascapes are
threatened by global climate change. Thus, to address this problem, it is essential
to understand the complex ecological relationships between the components of
coastal seascape systems and their links to land ecosystems, including the positive
and negative effects of humans. The physical and ecological relationships
between tropical landscapes and seascapes often dene the energy and matter
uxes through which human activities exert their inuence on coastal ecosys-
tems. We illustrate these relationships by presenting a case study and conceptual
model of coastal and tourism development in the Yucatan Peninsula, Mexico.
Scientic knowledge and proposals for better governance practices are available
to guide management actions and to protect the connectivity of seascape systems.
Nonetheless, political will, coupled with the implementation of local and regional
integrated management schemes of coastal zones, is urgent. More examples are
needed of inland and coastal human societies that successfully integrate scientic
knowledge of the links among ecosystems and their decision-making processes in
order to achieve sustainable development based on the services provided by their
surrounding seascapes.
Keywords
Tropical seascapes Connectivity Mangroves Seagrass Coral reefs Yucatn
Peninsula
1 Introduction
In tropical waters, seascapes, or interlinked patches of marine ecosystems ranging

from mangroves to seagrass beds to coral reefs constitute a major feature of the
coastal zone and are often located adjacent to each other (Ogden 1997). The tropical
seascape is a spatially heterogeneous area of coastal environment that can be
perceived as a mosaic of patches, a spatial gradient, or some other kind of geometric
pattern found in either benthic or pelagic environments (Bstrom et al. 2011). A
seascape encompasses meta-ecosystems, i.e., sets of ecosystems connected by
spatial ows of energy, materials, and organisms across ecosystem boundaries
(Loreau et al. 2003). This concept represents an expansion of the notion of meta-
communities through the integration of mass and energy transfers among these
ecosystems. It provides a theoretical framework for the understanding of interlinked
The Animal Forest and Its Socio-ecological Connections to Land and Coastal. . . 3
processes among different ecosystems, such as dissolved and particulate matter

uxes, the exchange of living material, the export and import of matter and energy,
and diversity-production patterns, among others (Loreau et al. 2003).
The capacity for the development of each particular ecosystem within a tropical
seascape can be enhanced by its interaction with other ecosystems. This may be one
of the reasons why tropical seascapes are major providers of natural resources and
ecosystem services for millions of human beings (Moberg and Rnnbk 2003). The
structural and functional values of the different mosaics of ecosystems that constitute
these tropical seascapes and their interactions with humans are highly relevant.
Interactions between seascape components (including the human societies living in
these areas) can largely be subdivided into chemical, biological, and physical uxes
(Ogden 1997). Examples of some of the interactions among ecosystems include
exchanges of larvae, juvenile and adult fauna, plankton and nutrients (Lowe and
Falter 2015), detritus, water, sediments (Nagelkerken 2009a), and sewage pollution
from terrestrial runoff (Wear and Vega-Thurber 2015).
1.1 Connectivity in Tropical Seascapes
The coastal tropical seascape is a continuum that spans inland ecosystems, coastal
ecosystems, and the ocean. Although rivers represent only a quarter of the coastal
area of the world, the vast majority of water and sediment is discharged through this
land-sea connection, so the ow and exchange of dissolved and particulate materials
from land to sea are substantial. Also, groundwater is an important source of
nutrients such as dissolved inorganic nitrogen (DIN) for coastal ecosystems and
coral reefs, particularly in areas with high coastal development. Submarine ground-
water discharge (SGD) brings an estimated range of 3800 mmol h1 of nitrogen per
meter of shoreline (Paytan et al. 2005). In areas with permeable and porous carbon-
ate karst geology, such as Yucatan, Mexico, the water nutrient quality is regulated by
groundwater discharges and may be similar in magnitude to that of rivers; these
inputs are linked to agro-industries, atmospheric deposition, urban-public activities,
and industrial activities, with an estimated nitrogen to phosphorus (N:P) discharge
ratio of 194.9:1 (Aranda-Cirerol et al. 2006).
Bouillon and Connolly (2009) consider that tropical rivers in coastal margins
deliver most of the carbon and nutrients supplied from land to sea. According to
Alongi et al. (2014), global water discharge from tropical rivers is higher in America
than in other continents, while the highest global sediment discharge is in Asia.
Tropical rivers channel up to 5564 % of the total global ux of riverine dissolved
inorganic carbon (DIC), with the highest yields taking place in Asia, where carbon-
ate rock is most abundant. Approximately 30 % of the global particulate inorganic
carbon (PIC) ux is discharged from the tropics. Equatorial rivers have lower DIC
but higher dissolved organic carbon (DOC) concentrations than those at other
latitudes. This can be attributed to the carbon-rich African soils, and the prevalence
of peat and black-water rivers in tropical Asia and the Americas. The total DOC
discharge accounts for 6366 % of the total global ux. Moreover, most of the
tropical particulate organic carbon (POC) discharge occurs in the Americas and
Asia, with the total POC discharge from the tropics estimated at 6470 % of the
global ux.
Oceans also shape the structure and function of tropical seascapes. For example,
currents carry out an enormous amount of larvae to tropical seascapes, where they
can further develop. Oceans are the main regional connection drivers among meta-
ecosystems, through waves, streams, coastal upwelling, and cyclones (Lowe and
Falter 2015). These physical processes transport key materials, nutrients, and plank-
ton, supporting life in the tropical seascapes. As Lowe and Falter (2015) mention,
although oceans play a fundamental role in shaping the distribution of species, a
modern understanding of the complex interactions between oceans and coral reefs is
emerging. Nevertheless, oceans play a critical role within the patterns of biodiversity
distribution through the dynamics of currents at different scales and the interaction of
nutrients among ecosystems. Oceans not only have an enormous inuence on the
growth, metabolism, and community structure (Hammer and Wolansky 1988) of the
different assemblages and ecosystems that compose a tropical seascape but also on
the hydrodynamic processes related to heat dissipation, transport and mass transfers
(nutrients and carbon), heat, temperature, and momentum (drag and dissipation)
(Lowe and Falter 2015).
Several tropical seascape features can contribute to the stability of local ecosys-
tems. These features include tropical forests, mangroves, seagrass, fringing reefs,
barrier reefs, the generation of continental margins, reef lagoons, and the control of
coastal erosion through the mitigation of energy from waves. Different aspects of the
tropical seascape structure and functioning depend on the geomorphology of the
coastal zone and continental shelf, which are connected through the movement of
inorganic matter, detritus, and living material (Loreau et al. 2003). In these sea-
scapes, mangroves and seagrass beds promote the development of offshore coral
reefs by interfering with discharges of freshwater and acting as sinks for pollutants
and organic materials, ltering the runoff water that may eventually reach coral reefs
(Harborne et al. 2006; Berkstrm et al. 2012; Gillis et al. 2014). These coastal
vegetation systems control the concentration of nutrients such as nitrogen and
phosphorus as well as suspended sediment in the water column. Nevertheless,
these ecosystems can also naturally be net exporters of nutrients to coral reefs
(Bouillon and Connolly 2009). In turn, coral reefs create a protective environment
for seagrass beds and mangroves by buffering the energy from oceanic waves and
currents (Harborne et al. 2006; Berkstrm et al. 2012) and by protecting the coast
from pounding waves, storms, and hurricanes (Gillis et al. 2014).
There are also movements of oating living material, such as macroalgae (for
instance, the brown alga Sargassum) and seagrass, which can strongly inuence
populations, consumer-resources, food availability, and community structure (Polis
et al. 1997). Biological interactions through the biological transfer of material among
different ecosystems due to ontogenetic migration or trophic relay (Bouillon and
Connolly 2009), as well as the transport of larvae from one ecosystem to another
may inuence the shape of the coral reef community, as in the case of sh assem-
blages (Mumby et al. 2004). It is known, for example, that both mangroves and
Fig. 1 The main climate change threats and impacts on the tropical seascape include: (1) ocean
warming, (2) ocean acidication, (3) increase in the frequency of tropical storms and hurricanes,
(4) sea-level rise, (5) decreasing dissolved oxygen concentrations (hypoxia), and (6) change in
rainfall patterns
seagrasses act as nursery grounds for sh recruits, decapods, and many other
organisms (Nagelkerken 2009a) or sh, which subsequently integrate into or interact
with coral reef communities (Nagelkerken 2009b; Mumby et al. 2004)
1.2 Global Climate Change Impacts on Animal Forests
The coastal tropical seascape is crucial for global climate change, particularly for the
circulation of heat, salinity, and water vapor (Alongi et al. 2013). While some
ecosystems are likely to benet from climate variations, coral reefs are extremely
vulnerable. For instance, coral bleaching can be caused by a gradual increase in
water temperature (12 C above the maximum temperature recorded), causing
stress and the expulsion of the endosymbiotic zooxanthellae that provide its main
energy source, as this bleaching causes their death (Munday et al. 2008). It is
estimated that over 16 % of coral reefs worldwide have suffered serious damage
from massive bleaching events (Wilson et al. 2006). Projections of temperature
models indicate that bleaching may become a recurring phenomenon worldwide in
the course of this century, threatening the ecological balance of seascape systems
(Buddemeier et al. 2011) (Fig. 1). In addition, 3350 % of coral reefs have been
largely or completely degraded by a combination of local factors and global climate
change (Wilkinson 2008).
The Fifth Assessment Report of the Intergovernmental Panel on Climate Change
(IPCC) provides high certainty that anthropogenic emissions have caused
measurable changes in the physico-chymical aspects of ocean environments, includ-

ing warming and associated phenomena such as sea-level rise, hypoxia, and acidi-
cation (Howes et al. 2015). These changes imply direct effects on ocean biological
processes. In the ocean, key biological processes such as the dispersal and replen-
ishment of larvae may undergo changes in association with the high sea surface
temperatures of the sea (Ayre and Hughes 2004). Increased ocean temperatures are
expected to affect larval development in corals, which, in turn, may lead to declines
in adult stocks and change temporary patterns of dispersal and gene ow. This may
eventually cause spatial and latitudinal constrictions or expansions (Ayre and
Hughes 2004) in populations. In fact, it has recently been suggested that high
temperature may lead to increases in the local retention of coral reef larvae (Figueiredo
et al. 2014).
Species population and their connectivity could also be compromised by the
increased fragmentation of reef habitats resulting from ocean acidication and
bleaching of coral reefs (Munday et al. 2008). Ocean acidication will cause
deciencies in calcication rates and a reduction of recruitment by disrupting coral
larval settlement and algal interactions, among other effects (Doropoulos
et al. 2012). It is estimated that drops in pH values by the formation of bicarbonate
ions (HCO3) in seawater due to surplus atmospheric carbon dioxide (CO2) may
reduce the availability of carbonate ions in seawater by up to 50 % by the year 2065
(Cao and Caldeira 2008). Carbonate ions are essential for organisms that produce
shells or exoskeletons based on calcium carbonate (CaCO3), such as corals, mol-
lusks, echinoderms, foraminifera, and crustaceans (Doropoulos et al. 2012).
Another impact associated with temperature is the concentration of dissolved
oxygen: water warming causes a decrease in the availability of oxygen (O2) in the
ocean (hypoxia) (Vaquer-Sunyer and Duarte 2008). Future scenarios of the concen-
tration of dissolved oxygen are not encouraging, as it has been predicted to decline
by 17 % worldwide in the next 100 years (Keeling et al. 2010). Most affected are
aquatic sh and invertebrates because their physiological mechanisms are deter-
mined by the concentration of dissolved oxygen in the water. The sensitivity and
susceptibility to hypoxia vary considerably among taxa (Vaquer-Sunyer and Duarte
2008). Thus, the biological and ecological impacts of hypoxia are extremely com-
plex and difcult to estimate (Prtner and Knust 2007).
Sea-level rise is perhaps the most pervasive threat to the integrity of seascapes
that encompass coastal lagoons, mangroves, and coral reefs. Higher sea levels may
result in a larger inuence of seawater in coastal systems where the balance between
salt and fresh water is critical to many ecological processes. Among these systems
are coastal lagoons, seagrass communities, and mangrove forests whose suitability
as nurseries for pelagic and reef species may be compromised by imbalances in
salinity (Harborne 2013). Indeed, Gilman et al. (2008) suggest that sea-level rise
may cause large-scale loss of mangrove habitat, coastal dunes, and lagoon ecosys-
tems. Sea-level rises may also result in increased wave energy and modications of
wave regimes that can fundamentally alter the morphodynamics of coasts. In the
case of reefs, the loss of coral crests due to bleaching and other processes may be
exacerbated by projected increases in annual mean wave heights and changes in
wave direction (Hemer et al. 2013). The loss of this wave mitigation effect may, in
turn, affect other physical and biological elements of reefs themselves and, in the
case of barrier reefs, even those of ecosystems located on the adjacent coast.
However, it has also been suggested that under sea-level rise scenarios, coral
cover, and reef growth may signicantly increase (Hopley 2011).
Coastal cities and small island nations are the most vulnerable to sea-level rise.
Social-ecological ecosystems are currently under threat and will further face severe
challenges due to global climate change. This may have unprecedented effects on
local communities that live on the coast and depend on shing and marine resource
extraction. It can impact nearly all sheries, coastal societies, and economies world-
wide because it may affect the biodiversity of seascapes as a whole.
2 Linkages and Complexities of Social and Ecological

Systems
Coastal ecosystems are transformed by humans and are facing a great variety of local
and global stressors that produce important changes for marine animal forests (Burke
et al. 2011). Within seascapes, coral reefs are among the worlds most diverse
ecosystems. A recent estimation establishes that coral reefs offer food and shelter
to 830,000 species (95 % condence interval: 550,0001,330,000 species) (Fisher
et al. 2015). Associated with this high diversity, coral reefs provide many valuable
services for humans, including shoreline protection, livelihoods from ecotourism,
shery production, and a living factory of compounds for biomedical use with a
global value of over US$ 31 billion (estimated for 2014) for all reefs combined
(Wear and Vega- Thurber 2015). Nevertheless, 75 % of coral reefs are currently at
risk from local stressors and thermal stress. It is envisaged that by 2030, over 90 % of
reefs might be at risk, along with the benets they provide (Jackson et al. 2014).
The great majority of global coral reefs are close to coastal zones. Globally,
human inuences are producing dramatic shifts from coral to algae cover and are
disrupting biodiversity (Acosta-Gonzlez et al. 2013) and ecosystem functioning
and services (Morberg and Folke 1999). A small disruption in biodiversity can cause
the collapse of key services that coral reefs provide, such as habitat for the recruit-
ment of commercial species. In turn, the loss of coral reef services may negatively
affect the resilience of the socio-ecological system as an imbalance develops
between human demand for such services and the actual capacity of ecosystems to
supply them.
Linking social and ecological systems is necessary to better understand the
complex interactions between humans and coral reefs. There is growing interest in
the dynamics and sustainability of the natural environment and human interactions
(Diaz et al. 2011) and the conceptualization of social-ecological systems (Biggs
et al. 2015). The socio-economic system can be characterized by the human activ-
ities that have been developed, such as tourism, urban development, or shing.
These drivers can impact tropical seascapes (seagrass, mangroves, and dunes) and
therefore lead to negative feedback loops in the ecosystems services. Nevertheless,
Fig. 2 A conceptual model demonstrating key processes, impacts, and negative feedbacks (yellow
arrows) within tropical coastal seascapes and the connectivity of land (brown arrows), mangroves
(green arrows), seagrass (blue arrows), and coral reefs (red arrows) in the Yucatan Peninsula,
Mexico (arrows/ows adapted from Silvestri and Kershaw 2010)
the management of seascape connectivity for instance, through the establishment of

marine protected areas or the protection of habitats may involve direct actions and
solutions to environmental problems and may include economic, political, and
cultural decisions that create positive feedback loops between the ecological and
socio-economic systems. Overall, the ecological system may be characterized by the
state of ecosystem services, such as herbivorous sh biomass that can control
macroalgae, the condition and architecture of coral reefs, commercial sh stocks,
sh biodiversity, and sh feeders that can control lionsh.
2.1 Transformations of the Yucatan Peninsula
2.1.1 Conceptual Model of the Yucatn Peninsula

In this section, we dene a conceptual model of the cross-interactions and connec-
tivity within seascapes, with a focus on the Yucatn Peninsula as a case study
(Fig. 2). In the model, the orange arrows show the negative feedbacks and the
impacts of the destruction of habitats such as mangrove vegetation, dunes, beaches,

and seagrass ecosystems and their inuences on the loss of coral cover and increased
macroalgae, leading to phase shifts in coral reefs due to increased nutrient inputs and
sedimentation. All these feedbacks result in a decrease of ecosystem services and
functional diversity, which may entail socio-economic changes for coastal
populations.
The main ecosystem functions provided by the connectivity of ecosystems are
also represented by colored arrows and are further described. The land (brown
arrows) inuences other habitats through sediments, nutrients, and freshwater dis-
charge, but also through pollutants carried through groundwater. Mangrove ecosys-
tems are the rst buffers to receive land inputs, which then help slow down the
freshwater ow; and along with sea grasses, both assist in the binding of sediment
loads and in the absorption of inorganic nutrients.
These nutrients catalyze primary production within the system and result in an
export of organic material for near shore and offshore webs. It is important to note
that the state of these interactions is related to the amount of nutrients in the system,
biogeochemical cycles, the balance/unbalance of the food web in each of the
ecosystems, and the overall functional diversity of species in the tropical seascape.
Overall, the seascape shares key services, such as storm buffering and the net export
of sh invertebrates in all life cycles (larval, juvenile, adults), areas of reproduction,
and habitat zones.
One example is the rapid transformation of the Caribbean coastal zone of the
northern Yucatn Peninsula (State of Quintana Roo, Mexico). The development of
Cancn and the Mayan Riviera from the 1970s to 1990s has caused profound
changes in the coastal seascape and the socio-ecological system (Torres and Monsen
2005) (Fig. 3). Cancn is by far one of the most dramatic cases of coastal transfor-
mation from a natural habitat to an urbanized tourism destination and is a picture-
perfect case for demonstrating how the alterations of a coastal ecosystem have
occurred over a short timescale (20 years) and continue to progress.
Cancn is the main beach tourism in Mexico, with visitors from all over the world
(4,013,032/year) and signicant revenues of $4,135 million dollars (USD) per year
(as of November 2014) (McCoy 2015). Despite the substantial economic benets
(jobs, income generation, international tourism destination) of this activity, Cancn
is far from being sustainable and is considered to have reached its environmental
capacity, with complex deteriorations in its social and cultural capital (McCoy
2015).
Moreover, multiple stressors act on the transformation of coastal seascapes on the
Caribbean coast of the Peninsula, including direct clearing of tropical forests and
mangroves, and the construction of housing developments for low-income families,
and the construction of hotels and other infrastructure on coastal dunes. Due to this
development, this area suffers severe problems with coastal erosion (Gonzlez-Leija
et al. 2013). Moreover, thousands of hotel rooms have been built, and many of the
low-income housing developments dispose of their sewage through septic tanks that
are connected to absorption wells (Torres and Monsen 2005). Even when sewage is
treated, only a fraction of it goes through water treatment plants. Hence, wastewater
Fig. 3 Urban transformation and growth in Cancn, northern Yucatan Peninsula, Mexico, due to
tourism expansion promoted by the Mexican Government, along with national and foreign inves-
tors. Satellite images taken on February 5, 1979(left) and January 24, 2009 (right). (Image source:
United Nations Environment Programme (UNEP), from Latin America and the Caribbean Atlas of
our Changing Environment, 2010)
and its pollutants eventually reach the groundwater and make their way into coastal
lagoons and reef areas (Hernndez-Terrones et al. 2015).
2.1.2 Geological Formations Influence the Tropical Seascape

The vulnerability of tropical seascapes in the Yucatn Peninsula is intimately related
to the geological dynamics of the region. The peninsula is a partially emergent
calcium carbonate platform extending to the north and northwest into a massive
continental shelf (Perry et al. 2003). There is almost no surface runoff on most of the
peninsula, and the aquifer system has developed within a nearly horizontal perme-
able limestone and dolostone karst whose oldest rocks are from the Tertiary period
(Perry et al. 2003). Thus, a fresh groundwater lens ows through this karstic plain
above a saline intrusion layer. The depth of the nearly at water table (gradient =
2 cm/km) is determined by elevation above mean sea level, the depth of the interface
between fresh and seawater, and, to a lesser extent, by the recharge of the aquifer
from annual precipitation, which increases from 500 to 1,500 mm along a northwest-
southeast axis. The water table is frequently exposed in places where the ceiling of
underground cavities has collapsed, forming sinkholes, or cenotes (from dzonot, the
Mayan word for well) as they are locally known, while in coastal areas, groundwater
usually forms springs (especially on the West and North coasts of the peninsula).
Other interactions include the ows of fresh ground water from karstic cave systems
into the sea, which are more often on the Caribbean coast. Overall, the balance of
fresh and seawater has been a major determinant of coastal geomorphology,
inuencing the formation of coastal lagoons and even coral reefs.
Another element that shapes coasts on the Yucatn Peninsula is the transport,
removal, and deposition of ne biogenic sand by currents and winds. This is the
predominant geomorphological process along the Northeast, North, and West coasts
of the peninsula. This constant transport of sand makes for a more turbid water
column than on the Caribbean coast of the Peninsula, and its clearest effect is the
formation of sand barrier islands on the Northeast, North, and West coasts, as
currents and waves coming inshore from the northeast transport and deposit sands
in an east-to-west direction. Thus, cul-de-sac lagoon systems eventually form, in
which the windward edge of the barrier island joins the continent by sand deposition,
while the leeward edge remains open. As a result, areas near the mouth of these
lagoons are under direct inuence of the sea, while conditions at the closed end are
more inuenced by evaporation, rainfall, and groundwater sources. This creates
salinity and nutrient gradients that result in a wide variety of terrestrial and aquatic
habitats. As the geomorphological development of these lagoons progresses, the
leeward mouth is nally blocked by sand, and they turn into enclosed lagoons and
wetland systems, where the balance among evaporation, rainfall, and groundwater
input becomes critical in dening ecological conditions.
The Caribbean coast of the Peninsula is substantially different from the others in
that karstic rock often reaches the sea, creating long extensions of rocky coast
interspersed with stretches of sandy beaches. Another major difference is the
absence of a vast continental shelf and the presence of the Mesoamerican Reef
(MAR), the worlds second-largest barrier coral reef, running parallel to the coast.
The absence of an extended continental shelf and the fact that sands are usually
conned to certain areas of the coast result in much clearer waters than on the North
and West coasts of the Peninsula. Groundwater usually reaches the coast and the sea
more directly than on the other coasts through karstic cave systems that were above
sea level during the last glacial period but have been ooded by seawater to different
degrees ever since that period ended.
3 Human-Derived Fluxes and Animal Forest Habitat Change
In addition to the complex impacts of global stressors, coastal ecosystems are

exposed to a number of problems associated with changes in the local environments.
Overall, these changes derive from alterations of chemical, biological, or physical
uxes, which are caused by the growth of human settlements and infrastructure,
habitat loss, resource overexploitation, and land reclamation activities that can
completely destroy whole components of tropical seascapes (Fabricius 2005).
As a complement to the conceptual model of a socio-economic and ecological
system, Figure 4 shows real case images of the complex direct and indirect interac-
tions. It also demonstrates the linkages of human-derived uxes (drivers, impacts,
and ecosystem state) that result in the loss of ecosystem services. The main drivers of
the socio-economic system (red) are related to tourism and urban development,
increases in migration and low-income housing, and overshing. The loss of key
habitats in the tropical seascape is shown in light brown. Such losses may cause
Socio-economic System
Tourism & urban Housing & Cruise ship

Overshing
development migraon industry
Mangrove loss Sink-hole polluon Dune loss Sea grass damage

Ecological System
Changes in
Euent & nutrients Disease & bleaching Phase shis funconal diversity
Loss of ecosystem services
Fig. 4 Interactions of complex drivers of change, impacts and ecosystem state within the socio-
ecological system of a tropical seascape (Image sources: INEGI, 2015; DigitalGlobe, Google Earth.
Elaboration: CINVESTAV-LEEAC, Mexico, and all other images: Andrea Rivera-Sosa)
coastal transformations to the ecological system (light blue). As mentioned before,

the ecosystem state (light blue) may be in distress due to phase shifts and decreased
resilience, whereas coral reefs may be exposed and prone to further impacts of
bleaching and disease outbreaks.
3.1 Chemical Fluxes
3.1.1 Water Quality and Nutrient Inputs

Locally, one of the most pressing issues is the decline in water quality in coastal
areas, which is associated with land transformations in order make way for agricul-
tural expansion and urban development. These changes alter watershed runoff and
chemical uxes from land to sea. Water quality parameters such as dissolved
inorganic nutrients (nitrogen and phosphorus) and suspended particulate organic
matter concentrations may increase due to these land use conversions and affect
coastal ecosystems by increasing turbidity, sedimentation, and pollution (Fabricius
2005).
Each of these parameters is associated with a medium to moderate negative

impact on calcication, tissue thickness, zooxanthellae density, photosynthesis,
and survival of adult colonies in reefs. In contrast, it has been documented that
particulate organic matter in the water column may have positive effects on the
ecological processes of coral reefs (Fabricius 2005). Corals can adapt to reduced
light caused by high turbidity in the water column by increasing lipid reserves in
polyps, thus ensuring energy levels necessary to survive low light conditions (May-
nard et al. 2010; Fabricius 2005). However, despite their ability to adapt to high
turbidity or sedimentation, corals are highly vulnerable to chemical toxins (heavy
metals, insecticides, and fungicides) that can be deleterious at different life history
stages, ranging from fertilization to larval development processes (Markey
et al. 2007).
Sewage pollution is often identied as a problem for coral reefs, but it has not
received the serious attention that is needed. As Wear and Vega-Thurber (2015)
recently noted, even treated sewage can introduce a variety of substressors to coral
reefs. These individual stressors chemically interact with one another, thus having
more intense negative effects on corals (e.g., nutrient-enhanced pathogen growth).
These synergistic interactions are poorly understood, and it is suspected that they
may potentiate the impacts of other nonrelated stressors, such as bleaching or
overshing leading to phase shifts. Redding et al. (2013) also showed an important
correlation between an increase in derived nitrogen from sewage pollution and
disease severity in the coral genus of Porites in Guam. Similarly, Mora (2008) and
Bozec et al. (2008) showed that in the Caribbean, the abundance of macroalgae and
coral mortality was correlated with human density in areas neighboring coral reefs.
Studies in the Pacic and Atlantic oceans are showing that phase shifts involve
not only transitions from coral to algal dominance but also from coral to sponge
dominance (McMurray et al. 2015). In particular, Knapp et al. (2013) suggested that
sponge dominance in the lagoon of the Palmyra Atoll, in the Central Pacic Ocean,
was a result of a decrease in environmental quality: sponges thrived as sediments,
turbidity, and food availability increased in an environment that was originally
dominated by Acropora species, crustose coralline algae, turf algae, and the giant
clam Tridacna maxima (Knapp et al. 2013). In another example, surveys conducted
in Florida between 2000 and 2012 showed that the population and recruitment of the
barrel sponge Xestospongia muta has been increasing over time, while scleractinian
coral cover has been decreasing (McMurray et al. 2015). In the Caribbean, major
phase shifts are causing changes in reef structure, as larger more structurally
complex species, such as those in the Orbicella complex, are being replaced by
less complex species, such as those in the genus Porites (Alvarez-Filip et al. 2013).
Excessive nutrient inputs not only cause phase shifts in coral reef ecosystems but
may also affect seagrass ecosystems (Fig. 5). The nitrogen and phosphorus derived
from ltration of septic tanks from coastal housing may cause the proliferation of
algae and cause direct competition for light, space, and nutrients in seagrass. This
problem may be exacerbated in lagoon systems where less nutrient dispersion occurs
due to high resident times of seawater due to the protection of barrier reef systems.
Fig. 5 Seagrass ecosystem

dominated by Thalassia
testudinum and Syringodium
liforme, covered by
lamentous green macroalgae
(Chaetomorphalinum) in Big
Corn Island, Nicaragua
(Image source: Andrea
Rivera-Sosa, Date: 09/3/2016)
3.2 Biological Fluxes
3.2.1 Phase Shifts

Stressors such as nutrients from sewage and groundwater pollution may be the main
precursors of phase shifts from coral-dominated to algae-dominated reefs although
they may act in conjunction with shing pressures, bleaching, and hurricane events
(Bozec et al. 2008). Contrary to what is generally proposed in the coral reef literature
(Burke et al. 2011), groundwater pollution on the Caribbean coast seems to have
resulted in a benthic phase-shift scenario that is thought to be independent of the
distribution of herbivorous sh (Bozec et al. 2008). The nutrient-rich wastewater
may itself cause the proliferation of macroalgae over a threshold where herbivores
may not be able to control them, even if herbivore populations are not affected
(Williams and Polunin 2001). This bottom-up process of the benthic phase shift has
seldom been documented, but it constitutes an interesting explanatory alternative to
the paradigm of shing reducing the abundance of herbivorous sh (mainly parrot
sh species) and allowing macroalgae to overgrow and outcompete corals (Russ
et al. 2015).
Recent work (e.g., Bozec et al. 2008; Kramer et al. 2015; Russ et al. 2015; Wear
and Vega-Thurber 2015) suggests that such a bottom-up control is relatively simpler
that the top-down control, as it only implies a change in habitat consisting of
increased nutrient availability (Russ et al. 2015). Thus, the underlying cause of
habitat change could also be from different sources of local or regional coastal
impacts and global climate change, such as dredging, sedimentation, sewage pollu-
tion, hurricanes, or coral bleaching. In contrast, top-down control may imply more
processes (Russ et al. 2015): (1) shing to reduce herbivore (i.e., parrot sh)
abundance, (2) reducing grazing pressure, (3) increasing cover of macroalgae, and
(4) decreasing coral cover (algae outcompetes adult corals for space or inhibits coral
recruitment).
Fig. 6 Phase shifts from a coral- to an algal-dominated coral reef in Mahahual reef, Mexico (Image
source: Roberto Hernndez-Landa, Date: 2010)
Particularly in the Mesoamerican reef region (MAR), different bottom-up pro-

cesses may produce benthic phase shifts, both at the local (Bozec et al. 2008) and
regional scales, even within marine protected areas where there has been an evident
increase of herbivorous sh (Kramer et al. 2015). This pattern is consistent with
those observed in coral reefs in the Philippines (Russ et al. 2015) and New Caledonia
(Carassou et al. 2013).
Moreover, although there are no quantications of the loss of species produced by
changes at a large scale, there has been a recent phase shift from coral to algae in the
Mahahual reef system, Quintana Roo, Mexico (Fig. 6). This area has shown that the
establishment of a cruise ship pier and the loss of vegetation cover linked to tourism
and housing expansion may have produced the loss of 40 sh species and 43 ha
(10 km2) of coral cover (Acosta-Gonzlez et al. 2013; Martnez-Rendis
et al. 2015). This type of massive tourism development is seen in many locations
in the Mesoamerican Region and may be another precursor of phase shifts.
3.2.2 Loss of Tropical Wetlands

Humans vastly benet from the ecology of mangrove ecosystems via provisioning
services (food and natural resources), regulating services (ecosystem processes), and
cultural (nonmaterial) services. Mangroves play a critical ecological role in provid-
ing breeding and nursery habitats for valuable commercial species and sheries
(Davidson 2014). Mangrove wetlands and seagrass habitats make crucial contribu-
tions in terms of biogeochemical processes, serving as sinks for inorganic nutrients
and sediments and as exporters of organic matter (Childers et al. 1999). However,
changes in temperature, salinity, and extensive development are also leading to
major decreases in seagrass ecosystems, which through ecological connections can
negatively affect other components of the seascape.
Despite their vast ecological services, tropical wetlands are among the most
vulnerable ecosystems and are disappearing at an alarming rate. An estimated
5457 % of mangrove ecosystems have been lost over the last century (Davidson
2014). Wetland cover losses are linked to the overexploitation of natural resources,
modication and/or reclamation of land for urban development, and aquaculture
operations (Davidson 2014).
In Mexico, it is estimated that in the last 25 years, at least 10 % of the mangrove
ecosystems and cover have been lost or severely altered (Valderrama et al. 2014).
Many of the tropical wetland ecosystems in the Cancn area and Riviera Maya have
been vulnerable since the tourism expansion of the 1970s. Due to increased pressure
to preserve important habitats, a series of Protected Areas were designated. Even
though wetlands are under current protection, those located adjacent to or near the
areas of the urban expansion of Cancn continue to be affected.
The government, through the National Fund for Tourism Development (Spanish
acronym FONATUR) and other federal sectors, actively promotes tourism develop-
ment through the purchasing of land with tourism potential and the reselling of it to
interested investors (Saldivar 2016). In the years 2005, 2009, and 2011, the Mexican
Secretariat of Natural Resources (Spanish acronym SEMARNAT) provided legal
permission for the construction and expansion of new tourism developments for the
Tajamar Esplanade (74.24 ha) (SEMARNAT 2016). However, The General Wildlife
Law (Article 60-TER-1/02/2007) states, no removal, lling, transplanting, pruning
or any work or activity shall intervene in the integrity of the hydrologic ow of a
mangrove ecosystem (SEMARNAT 2016). Despite this, the federal government
gave permission for the destruction of 49.10 ha (total area varies depending on the
information source) of wetland ecosystem to expand the Tajamar Esplanade in
Cancn, causing a major national and international uproar from the civil society
and nonprot organizations (Fig. 7).
Overall, there are many issues with the interpretation of laws for the formal
protection of the wetlands and natural resources, yet political or economic interests
may inuence the decision-making processes. For instance, the FONATUR donated
to SEMARNAT a total of 3,533 ha to be designated a Natural Protected area, namely,
the Mangroves of Nichupt and Ecopark Cancn (107 ha-2013) (SEMARNAT
2016), but this donation may be used in strategic political decisions to allow the
development of certain areas. Moreover, in response to the environmental destruc-
tion, the federal government issued a public statement that mentioned: If permission
had not been given to clear cut the area, the lawsuits of the investors who acquired
the land with the permission of SERMANAT and FONATUR would have resulted in
a potential nancial loss to the Mexican government of more than three billion
Mexican pesos, which is equivalent to more than half the budget of the Mexican
Ministry of Tourism for 2016. Currently, the project has been temporally
suspended (SEMARNAT 2016; Saldivar 2016).
The destruction of wetland and mangrove ecosystems is an eminent threat to
coastal areas with high tourism demand. However, concerns and misconceptions
have arisen regarding the recovery and restoration of these ecosystems. Once the
ecosystems have been lost, the trajectories of recovery are extremely costly, and their
Fig. 7 Destruction of
mangrove and wetland
ecosystems of the
controversial project Tajamar
Esplanade, Mexico, due to
tourism development and
expansion (Image source:
Christian Alva-Basurto, Date:
14/03/2016)
success can only be measured in the long term. The urgent need to recover essential
wetland ecosystems has led to an emerging eld of restoration, which includes the
de-canalization of wetlands and rivers, rehabilitation of degraded oodplains,
decommissioning of dams, replanting of vegetation and wildlife, and the implemen-
tation of conservation and management plans to protect wetland ecosystems. This
has been partly due to the economic valuation of wetlands and the fact that more than
half of the worlds diverse wetlands have disappeared (Davidson 2014).
The eld of economic valuation of wetlands offers an opportunity to place a
monetary value on a wetland that reects its ecological and social importance in
order to ensure its protection. For this reason, scientists are increasing their under-
standing of the major carbon stocks and uxes in mangrove ecosystems as well as
how they vary across species and zones to determine how much organic carbon can
be stored (aboveground or at various layers underground). According to Herrera
et al. (2016), many regions in Mexico, particularly those in the Gulf of Mexico,
along with riverine type mangroves, contain species that can be considered high
carbon sink facilitators. Also, mangroves in the Yucatan Peninsula were found to
have the highest content of organic carbon (218.98 Mg C ha 1) in the rst layer of
underground substrate (030 cm), followed by mangroves of the Gulf of Mexico
Herrera et al. (2016). This important information allows for the estimation of total
captured carbon, providing data to quantify the total value of an ecosystem and its
loss when converted. This would enable the establishment of negotiating mecha-
nisms for the conservation of wetland habitats to prevent and discourage the
conversion of natural lands to lled lands for construction. In addition, this conver-
sion of land releases the carbon stored in the soil, increasing the carbon dioxide and
methane concentrations that contribute to climate change (Herrera et al. 2016)
Fig. 8 Interaction of the

constructed coastline and
dunes in Cancn, Mexico
(Image source: Andrea
Rivera-Sosa, Date: 30/06/
2009)
3.3 Physical Fluxes
3.3.1 Coastline Modifications

Tourism development continually demands a larger area of coastline, increasing the
interface where natural processes interact with built-up areas and coastal dunes
(Fig. 8). In general, there are certain natural and anthropogenic factors that inuence
coastal erosion on different timescales. The most important coastal processes affect-
ing sediment transport and the building up of coastal dunes are hydrodynamic factors
such as waves, currents, and tides, and wind (Masselink and Hughes 2003). Other
short-term events such as storms may also inuence coastal erosion and dune
formation.
Naturally, coastal dunes develop in areas where there is sufcient wind to
transport sediment (i.e., ne sand), after which vegetation growth stabilizes the
sand. Thus, vegetation plays a very important role in securing the sand dunes and
preventing wind erosion. Also, vegetation increases the fertility of the sand and
favors the accumulation of debris, which then helps moisture to be retained. Over
time, the dune increases in size, offering natural protection against beach erosion and
serving as a sediment storage system (Masselink and Hughes 2003).
These dynamic processes vary in different areas, but it is indisputable that human
settlements have a huge impact on beach recovery. Alteration of dune formation and
vegetation for the construction of roads and other urban infrastructure considerably
impacts the geomorphology of a coastal area (Fig. 9).
For instance, the roads not only have a direct impact on coastal dunes, but they
also allow a continual ow of trafc and people, which continues to alter dunes.
Sediment transport can also become unbalanced by modications of the coastline,
such as those caused by piers, groins, sea walls, boardwalks, and other human-made
structures (Masselink and Hughes 2003). Moreover, the direct removal of sensitive
components of ecosystems is widespread. As a result of coastal development, many
hotels and resorts remove seagrass to make the beach more suitable for swimming.
Fig. 9 Coastal roads built on

dunes primarily for tourism
transit on the island of
Cozumel, Mexico (Image
source: Moritz Kuch, Date:
24/11/2015)
Fig. 10 Coastal and marine

attractions for cruise ship
visitors concentrated at the
tropical seascape in Cozumel,
Mexico (Image source:
Andrea Rivera-Sosa, Date:
23/11/2015)
Excessive and intensive tourism-related aquatic activities and attractions are con-
centrated for cruise ship visitors, further altering seagrass and coastal dunes
(Fig. 10). Other impacts involve mechanical damage to corals or subaquatic vege-
tation due to recreational diving, anchoring, shing (nets), and marine transportation.
There are even more extreme cases of coastal changes. This is the case of the
Spratly Islands, south of China, where biodiverse atoll and seagrass ecosystems have
been completely dredged and built over to create functional islands so that the
government can gain control over territorial seas (Larson 2015). In addition, to
complete habitat destruction, land reclamation, and dredging can produce sediment
plumes and can reduce the availability of sediment, affecting the coastal morphology
and processes in distant parts of the seascape (Masselink and Hughes 2003). In this
process, the connectivity between populations of key species is also hampered
(Larson 2015).
4 Management and Conservation of the Animal Forest

and Tropical Seascapes
Worldwide, the main mechanism for the conservation and management of the
tropical seascape is known as the Marine Protected Area (MPA). It is estimated
that 27 % of all coral reefs worldwide are currently within MPA schemes, but only
6 % of these areas are considered effectively managed (Ct and Reynolds 2006;
Burke et al. 2011). In Caribbean countries, there are more than 630 MPAs,
encompassing 30 % of the regions coral reefs, and many locations still have to
increase their areas of protection (Burke et al. 2011). For instance, Mexico, one of
the largest countries in Latin America, has MPAs on 22.7 % of its territorial sea, on
12 % of its continental platform, and on 1.5 % of its Exclusive Economic Zone
(EEZ) (Bezaury-Creel et al. 2009).
MPAs are specically delimited geographical areas that cover extensions of the
continental platform and the seascape. MPAs can be internationally or nationally
declared for the conservation of biodiversity, resources, environmental services, and
cultural heritage (Kelleher 1999). Their main objective is the protection of habitats
from destructive activities, the recovery, and productivity of sheries, and the
implementation of actions to increase resilience to impacts and maintain ecological
processes across the seascape (Moberg and Folke 1999; Pandol et al. 2005; Mumby
2013). These areas may be established under a wide variety of criteria, standards,
and levels of protection, which may include marine spatial planning, no-take
reserves, regulation of sheries, and accepted levels of resource exploitation
(Aswani et al. 2015). Their design and operation might be federal and/or state-
based, comanaged (NGO and state-based or private entities and the state), top-down
(the most common), or bottom-up (community-based), run by indigenous organiza-
tions and/or include a mixture of involvement. The increasing number of different
comanagement schemes is a result of the lack of direct funding from state and a lack
of adequate management and implementation.
Efforts to preserve key marine ecosystems face major challenges due to a lack of
funding, low capacity, and the absence of political will to enforce the environmental
laws and regulations at the core of MPAs. This can effectively turn MPAs into paper
parks (Mora et al. 2006), where conicts with political interests (such as industrial
shing or tourism development), and problems with communities whose livelihoods
depend directly on scarce natural resources cannot be managed. Moreover, when the
capacities for management and effective intervention are lacking, scientic knowl-
edge may actually become irrelevant as technical documents and recommendations
are put forward but not implemented (Bearzi 2007).
Indeed, it is estimated that two-thirds of conservation assessments published in
peer-reviewed scientic literature are not followed by conservation actions (Bearzi
2007). Another common problem between conservation science and the real world
(Knight et al. 2008) is that few policy makers are willing to face the great challenge
of inspiring people to protect the environment. Marine conservation requires a long-
term commitment to pursue direct actions and a multidisciplinary approach to benet
marine ecosystems. In addition to solid science and well-conceived action plans,
what is desperately needed to promote MPAs is public pressure (Bearzi 2007). An

effective MPA plan is one that links knowing and doing (Knight et al. 2008).
4.1 Social Inclusion in the Management of MPAs
A complex issue in social-ecological systems is when people and/or indigenous

communities have been part of a seascape and have depended on its productivity
long before it is declared an ecologically sensitive area to be protected. Also, many
settlements within MPAs continue to live in poverty under precarious and marginal
conditions (Bezaury-Creel et al. 2009).
The establishment of MPAs and the inclusion of indigenous and local populations
to ensure effective and rights-based access to resources has been a growing eld of
study, with controversial cases throughout the world that need to be constantly
revisited. Case studies conducted globally have all argued the importance of the
inclusion of local people in the establishment, management, and enforcement of
MPAs (Christie et al. 2009). In many cases, livelihood alternatives are sought.
Eco-tourism and community-based tourism (CBT) also seem to be proposed often.
However, it is important to mention that the establishment of an MPA and its
management plan from a top-down approach may include programs for economic
alternatives for the community that may not always be welcomed and perceived as a
good initiative by local populations. Tensions, conicts, and nonapproval can result
from an improper process. Community-based tourism does not always work, and the
mechanisms of its employment (or the politics of the MPA) may entail a variety of
complex sociological and ecological factors that need to be taken into account.
For example, in Honduras, the rst MPA in the country (the Natural Marine
Archipelago Monument in Cayos Cochinos-1993) was promoted by national inves-
tors and a philanthropist, along with technical assistance from the Smithsonian
Institution and nancial support from international organizations. This area com-
prises 2 main islands and 13 cays with important seascape ecosystems, and it has a
long history of resistance from indigenous shing communities that benet from the
area (Brondo and Woods 2007). To foster acceptance among communities, the
Honduran Coral Reef Foundation (HCRF) (a comanaging authority), local NGOs,
and international organizations have promoted community-based tourism programs
(in their management plans since 2004) as an alternative to shing through training
and condence-building (Brondo and Woods 2007).
However, recent changes in the comanagement structure have provided a 10-year
concession to the Mexican Azteca Foundation, which has resulted in other priorities
related to tourism development (Kuch 2015). There are resource-based restrictions,
and the majorities of the cays are privatized and have limited local communities use
to one island and two cays. Moreover, the new concession actively promotes the
lming of international reality shows (which are seeing increasing revenues abroad)
in the sensitive habitats of the MPA (in cays designated for bird and turtle nesting
habitats within the management plan) that have closed access to local indigenous
people. This type of tourism confers very little benets upon the locals (750$ each
per 4 communities/per lming), and the management is further creating social

inequalities (Kuch 2015). This highlights the negative impacts of a nonsustainable
type of tourism on the socio-ecological system. This case is interesting because it
shows how neoliberalism can overrule the goals of sustainability included in the
management plans of an MPA, which ends up further disrupting indigenous groups
by creating double standards in favor of groups with higher capital (Brondo and
Woods 2007; Kuch 2015).
4.2 Current and Future Management Tools
Optimistic scenarios have resulted from the implementation of MPAs, particularly

shery management tools, which have restored overharvested stocks and helped
protect sh and invertebrate populations (MacNeil et al. 2015). Historically, the
exploitation of sheries (large vertebrates and shellsh) may have been the rst large
human disturbance of coastal ecosystems (Jackson et al. 2014). There is also
adequate evidence that shing affects food webs, population structures, and the
distribution of species and that it contributes to the loss of biodiversity and ecosys-
tem services. There is also evidence that the alteration of specic functional groups,
such as herbivores, leads to the cascading deterioration of reef communities (Jackson
et al. 2014).
For this reason, MPAs need a mixture of adaptive management schemes to work
alongside land use planning across all sectors (tourism, urban development) and the
implementation of marine spatial planning to enforce shing areas and recreational
areas (Fig. 11). Enforced zones that target no-take zones, closed seasons, and size
and species restrictions have demonstrated proven results in the restoration of food
webs from previously overshed conditions (Bellwood et al. 2004). Other actions in
MPAs involve sustainable tourism practices and the establishment of educational
centers to increase awareness.
In addition, a watershed- and ecosystem-based management approach is often
needed for the employment of measures such as reducing fertilizer inputs, increasing
forest cover, and installing sewage treatment plants to reduce sedimentation and
nutrient pollution (Burke et al. 2011). The realization that bottom-up processes may
be just as important as top-down processes in bringing about phase shifts in coral
reefs is extremely important for the management of seascape ecosystems. Preserving
coral-dominated reefs may require regulating not only the shing of herbivorous sh
but also the growth of human settlements on the coast, in addition to requiring the
treatment and disposal of wastewaters. Even if eutrophication and pollution threaten
coral reefs as much as overshing (Burke et al. 2011; Wear and Vega-Thurber 2015),
eutrophication has received considerably more attention from conservation organi-
zations, possibly because of the practical challenges of dealing with a large-scale
threat. For instance, pollution sources are often multiple, geographically widespread
or far from the reefs they affect. These factors, combined with the diversity of
pollutants involved, the high cost of water treatment systems, and the need to
Fig. 11 Management of tropical coastal seascape connectivity demonstrating the importance of

adaptive management schemes in Marine Protected Areas located adjacent to areas of high tourism
development
coordinate different government levels and jurisdictions, make management actions

difcult to implement.
Conservation initiatives in the MAR region have recently focused on
empowering MPA managers to understand elements of resilience and successfully
ameliorate the impact of local and global stressors on the regions reefs (Maynard
et al. 2010). A new review by Aswani et al. (2015) provides an excellent summary
and recommendations on how to move forward with emerging areas of science that
may offer tools for the development and improvement of management actions
regarding coral reefs. Overall, the main ndings show that new technologies in
molecular biology and marine biogeochemistry (stable isotopes) can be applied to
isolate cause-and-effect relationships between stressors and biotic responses. They
can also be used to determine the genotypic diversity within a population to further
understand the levels of connectivity among tropical seascape populations. In
addition, as global climate change continues to further impact ecosystems, scientists
are studying genetic intervention strategies in the eld of restoration by focusing on
thermo-tolerant and disease-resistant corals from local source colonies and the
translocation of these species to other areas. These actions must be followed to
enable the effectiveness of mitigation and management efforts to be evaluated in
order to provide insight into the potential resilience of particular coral reef species
and genotypes to both local and global stressors (Aswani et al. 2015). In addition, the
study promotes the shift in MPA designation and approach from the sole protection
of pristine locations to areas under continual stress, where species with built
resilience have been adapting over time and may offer genes tolerant to the tropical
seascape.
4.3 Governance to Improve the Management of Seascape

Connectivity
The creation of marine biological corridors (MBCs) has been widely encouraged as a
response to the vulnerable condition of seascapes (McCook et al. 2010). The concept
of MBCs was adapted from the Biological Corridor Program, a strategy to provide
connectivity between protected areas that was implemented in the Mesoamerican
region to enhance transcontinental biotic exchange, ensure gene ow, protect the
migration routes, and widen the protection of habitats (Ankersen 1994). The same
goals apply to MBCs, which consist of protected stretches of marine and coastal
ecosystems linking national and international MPAs. In its simplest form, a protected
corridor would be established to bridge a gap between two identical ecosystems
encompassed by two neighboring MPAs. For instance, two MPAs covering two
nonadjoining parts of a barrier reef could be linked by protecting the barrier reef
areas between them. However, a more integrated view of corridors could protect
areas linking landscapes and seascapes through places where crucial geochemical
uxes occur, currents ow, or migrations take place. The management of corridors
would thus focus on protecting key links between MPAs, sometimes regulating and
monitoring human activities that do not obviously or directly affect the MPAs.
Although the effectiveness of implementing MBCs has faced criticism, the concept
seems promising. Overall, the purposes of establishing conservation and manage-
ment programs for marine and coastal corridors are to protect key ecosystem
processes and strengthen the resilience of these ecosystems to disturbances and
uctuations (Maynard et al. 2010).
To holistically manage the complex interactions of the tropical seascape, govern-
ments, key actors, and stakeholders require the implementation of Integrated Coastal
Zone Management (ICZM) approaches. ICZM, together with watershed approaches,
aims at creating sustainable coastal development initiatives in which policy makers
can jointly protect resources while enhancing economic development. The ultimate
goal is to devise development strategies based on local protection and management
plans in order to reduce anthropogenic threats and propose overarching solutions. To
achieve integrated management, the following minimum criteria need to be met:
(1) a comprehensive vision of land use and planning policies; (2) implementation of
water and sanitation regulations; (3) MPAS and zoning of areas for public use
(including tourism); and, of course, (4) agreements and synergies among key actors
and stakeholders to move forward. In addition, adapted approaches such as marine
ecosystem-based management (EBM) in tropical regions accentuate the need to
include social systems and governance at their core (Christie et al. 2009). Moreover,
the level of success of ICZM or EBM schemes will depend on how policy is
implemented and adopted by all involved sectors and key actors. When smaller-
scale management has been complex, management over large geographic scales
such as the Caribbean region has proven to be extremely difcult.
In the Caribbean, the main industrial sheries have targeted ecologically impor-
tant species such as queen conch (Lobatus gigas), lobster, shrimp, groupers, and
snappers, all of which are in decline (WECAFC 2011). More recently, commercial
harvesting of sea cucumbers and historical catches of sharks, rays, and chimeras
have been recorded (WECAFC 2011). Moreover, shing pressures are simulta-
neously placed on coastal mangrove forests and coral reef systems, which enhance
negative feedback loops between these components of the seascape. It has been
observed that in many developing countries such as Honduras in Central America,
the lack of regulations or the means to enforce them can result in the use of
undersized nets that target juveniles in coastal lagoons and in large-scale shing
operations targeting spawning aggregation sites, further threatening sensitive and
endangered species such as groupers and snappers.
Despite this complexity, there are several success stories in the MAR region. For
example, the seven Central American countries enforced regional regulations for the
Caribbean spiny lobster (Panulirus argus). These regulations include size limits,
closed seasons, and export regulations (AECID/OSPESCA/SICA 2009).
Also, regional agreements and national and international environmental legisla-
tion have increased across all countries in the MAR. In particular, political interests
in the conservation of trans-boundary tropical seascapes (such as the declaration of
Tulum in 1997) (Kramer et al. 2015) have set an important path for the establishment
of The Convention for the Protection and Development of the Marine Environment
in the Wider Caribbean Region (WCR). The WCR includes very important technical
agreements such as the Protocols on Oil Spills, Specially Protected Areas and
Wildlife (SPAW), and Land-Based Sources of Marine Pollution (LBS). These
agreements have served as governmental and international platforms to implement
local strategies (Caribbean Environment Program 2015).
Leadership in organizations such as the Healthy Reefs Initiative, which coordi-
nates and gathers information with over 60 key actors in Mexico, Belize, Guatemala,
and Honduras, has been successful in promoting management actions based on
science in the MAR region (Kramer et al. 2015). In particular, actions in Belize to
establish moratoria on sheries targeting herbivores, as well as banned petroleum
exploration in MPAs and nearby zones, are a recent achievement of many institu-
tions working together. The rst shark sanctuary in the Americas has been
established in Honduras, representing the implementation of key political decisions
to preserve marine ecosystems. Moreover, in Quintana Roo, Mexico, there have
been initiatives to manage one of its largest protected areas, known as the Sian Kaan
Biosphere Reserve (6,510 km2), using an ecological conceptual model that is driven
by local and national societal needs and is implemented as an integrated resource
management plan (Mazzotti et al. 2005).
For these reasons, new cross-sector political agendas and creative solutions need
to be developed in order to ensure governance for the true management and
sustainability of the tropical coastal seascape. Aswani et al. (2015) recommend the
establishment of ve principles to ensure effective governance:
1. Develop leadership and raise awareness

2. Develop hybrid institutions or novel institutional collaborations (sustainable
funding mechanisms)
3. Establish clear and fair rules with conict resolution mechanisms
4. Experiment and adapt
5. Develop evaluation programs that monitor both social and ecological systems
Finally, Finkbeiner and Basurto (2015) recommend that societies rapidly adjust
from regular comanagement schemes to seeking multilevel comanagement and
peer-to-peer/governance networks. Doing so may require democratic participation in
the creation of hybrid comanagement systems that can be tailored to historical,
cultural, and ecological contexts.
5 Conclusions
Undoubtedly, achieving the much sought-after development of coastal human soci-

eties without compromising the animal forests and tropical seascapes of which they
are a key component will require a deep understanding of the complex socio-
ecological networks connecting the physical, chemical, biological, and human
components of such seascapes. Accepting that human activities and the transforma-
tions they entail are an intrinsic aspect of most modern ecosystems is fundamental to
establishing governance mechanisms and achieving sustainable interactions within
socio-ecological systems.
It is also critical to realize that human settlements located inland, away from
coastal areas, can have profound effects on the ecology of seascapes and the services
they provide to other human populations. These complex inter-relations need to be
documented, measured, and incorporated into models and management programs
over considerably larger geographical scales than those considered so far. If we are to
successfully tackle the challenge of sensibly using the resources and services that
coastal seascapes offer, we will need creative solutions based on sound scientic and
technical knowledge.
Scientists and coral reef managers now have the tools to understand the natural
resources, ecological processes, and value of ecosystem services that allow better
management decisions (Burke et al. 2011). Nevertheless, many of the actions needed
are highly dependent on the power of political will and commitment. It is important
to be aware that from a management point of view, knowledge alone can be of little
use without the involvement of all those stakeholders whose actions most directly
affect natural resources and services. The integrated management of social systems,
seascapes, and animal forests will inevitably require changing some of our practices
so that our negative impact on these natural systems is reduced and resilience
mechanisms can absorb it. Perhaps among the most difcult practices to modify
will be those derived from political and economic inertia among some of the most
powerful stakeholders. Again, an adequate regulatory framework, availability of
resources to enforce it, and political will are crucial. Both top-down (e.g., institu-
tional support, enforcement, and legislation) and bottom-up (proactive inclusion of
community governance systems) approaches need to be integrated (Aswani
et al. 2015). Overall, the management of the connectivity of the tropical seascape
needs to be adaptive, holistic, and efcient in order to decrease negative feedback
loops.
The full protection of remnant sensitive areas will never be achievable if govern-
ments continue to overlook their own environmental legislation and actively pro-
mote the destruction of natural habitats to fulll neoliberal agendas. Civil society has
an important duty to hold governments accountable. This is not an easy task, and
there still are many transnational obstacles to overcome, such as implementing
political decisions to reduce carbon emissions worldwide. There are also more
complex social factors that are likely to further inuence coastal seascapes in the
MAR region. For instance, in addition to traditional landscape transformations,
large-scale forest destruction and land conversion to pasture, cattle ranching, and
agriculture (oil palm) are now being nancially propelled by organized crime
(McSweeney et al. 2014). How such an increase in land use conversion will further
impact tropical seascapes needs to be explored in the near future. As mentioned
before, the overarching problems hindering the effective implementation of man-
agement, such as corruption, lack of governmental will, and economic interests that
hinder conservation, need to be tackled.
Finally, the understanding of socio-ecological systems needs to be coupled with
widespread involvement and commitment from society to realize a rapid increase in
success stories, showing that sustainable development based on the wealth of animal
forests is, in fact, possible.
6 Cross-References
Complexity and Biodiversity in Caribbean Coral Reefs

Conservation and Management of Vulnerable Marine Benthic Ecosystems
Ecosystem-Based Management: Opportunities and Challenges for Application in
the Ocean Forest
Resilience of the Marine Animal Forest
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The Animal Forest and Its Socio-ecological

Connections to Land and Coastal

Jess Ernesto Arias-Gonzlez, Andrea Rivera-Sosa, Jaime Zaldvar-

J.E. Arias-Gonzlez (*) A. Rivera-Sosa C. Corts-Useche

# Springer International Publishing Switzerland 2016 1

In tropical waters, seascapes, or interlinked patches of marine ecosystems ranging

processes among different ecosystems, such as dissolved and particulate matter

1.1 Connectivity in Tropical Seascapes

1.2 Global Climate Change Impacts on Animal Forests

measurable changes in the physico-chymical aspects of ocean environments, includ-

2 Linkages and Complexities of Social and Ecological

the management of seascape connectivity for instance, through the establishment of

2.1 Transformations of the Yucatan Peninsula

2.1.1 Conceptual Model of the Yucatn Peninsula

impacts of the destruction of habitats such as mangrove vegetation, dunes, beaches,

2.1.2 Geological Formations Influence the Tropical Seascape

3 Human-Derived Fluxes and Animal Forest Habitat Change

In addition to the complex impacts of global stressors, coastal ecosystems are

Tourism & urban Housing & Cruise ship

Mangrove loss Sink-hole polluon Dune loss Sea grass damage

Loss of ecosystem services

coastal transformations to the ecological system (light blue). As mentioned before,

3.1 Chemical Fluxes

3.1.1 Water Quality and Nutrient Inputs

Each of these parameters is associated with a medium to moderate negative

Fig. 5 Seagrass ecosystem

3.2 Biological Fluxes

3.2.1 Phase Shifts

Particularly in the Mesoamerican reef region (MAR), different bottom-up pro-

3.2.2 Loss of Tropical Wetlands

Fig. 8 Interaction of the

3.3 Physical Fluxes

3.3.1 Coastline Modifications

Fig. 9 Coastal roads built on

Fig. 10 Coastal and marine

4 Management and Conservation of the Animal Forest

what is desperately needed to promote MPAs is public pressure (Bearzi 2007). An

4.1 Social Inclusion in the Management of MPAs

A complex issue in social-ecological systems is when people and/or indigenous

per 4 communities/per lming), and the management is further creating social

4.2 Current and Future Management Tools

Optimistic scenarios have resulted from the implementation of MPAs, particularly

Fig. 11 Management of tropical coastal seascape connectivity demonstrating the importance of

coordinate different government levels and jurisdictions, make management actions

4.3 Governance to Improve the Management of Seascape

1. Develop leadership and raise awareness

Undoubtedly, achieving the much sought-after development of coastal human soci-

Complexity and Biodiversity in Caribbean Coral Reefs

Valderrama L, Troche C, Rodrguez MT, Mrquez D, Vzquez B, Velzquez S, Vzquez A, Cruz

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