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PREDICTIVE MODELLING OF WETLAND HABITATS IN
THE EBRO DELTA WITH A GIS APPROACH
 
 
 
 
Xavier Benito Granell
Màster en Planificació territorial: informació, eines i mètodes
Facultat de Turisme i Geografia
Universitat Rovira i Virgili
IRTA – Unitat d’Ecosistemes Aquàtics
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PREDICTIVE MODELLING OF WETLAND HABITATS IN THE EBRO DELTA
WITH A GIS APPROACH
Memòria del treball final del màster oficial de Planificació territorial: informació, eines i mètodes.
Per: Xavier Benito Granell
Dirigit per:
Dr. Carles Ibàñez Martí
Unitat d’Ecosistemes Aquàtics
IRTA – Sant Carles de la Ràpita
Dra. Rosa Trobajo Pujadas
Unitat d’Ecosistemes Aquàtics
IRTA – Sant Carles de la Ràpita
Dra. Yolanda Pérez Albert
Departament de Geografia
Universitat Rovira i Virgili
Vila-seca, Juliol de 2012
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Agraïments
Aquest treball ha estat possible gràcies una beca predoctoral de la Universitat Rovira i Virgili
dins del conveni URV-IRTA. La base cartogràfica (Model digital d’elevació i ortofotomapes)
són propietat de l’Institut Cartogràfic de Catalunya (www.icc.cat).
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Table of contents
Abstract ...................................................................................................................................... 5
1. Introduction............................................................................................................................ 6
1.1 General context: deltaic system, wetland habitats and human colonization .................... 6
1.2 The aquatic habitats of the Ebro Delta ............................................................................. 7
1.3 Predictive habitat modelling........................................................................................... 10
2. Hypotheses and objectives ................................................................................................... 15
3. Methods................................................................................................................................ 17
3.1 Study area ....................................................................................................................... 17
3.2 Wetland habitats, terrain variables and hydrologic alterations....................................... 19
3.3 Dependent variable: current distribution of wetland habitats......................................... 20
3.4 The independent variables: elevation and distances to hydrologic boundaries.............. 33
3.5 Vegetation transects........................................................................................................ 37
3.6 GIS development ............................................................................................................ 38
3.7 Statistical analysis........................................................................................................... 45
3.8 Model implementation in the GIS .................................................................................. 47
4. Results and discussion.......................................................................................................... 48
4.1 Current distribution of wetland habitats in the Ebro Delta............................................. 48
4.2 Logistic regression.......................................................................................................... 70
4.3 Probability of occurrence................................................................................................ 85
5. General conclusions ............................................................................................................. 91
6. References............................................................................................................................ 94
 
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Abstract
Predictive habitat distribution models, derived by combining multivariate statistical methods
with Geographical Information System (GIS) techniques, have been recognised for their
utility in ecological modelling. The knowledge about potential distribution of natural habitats
requires the link between current presence or absence of biological communities and a set of
relevant environmental variables. This work examines the feasibility of using multiple logistic
regression to model wetland habitat distribution in the Ebro Delta and to analyse how the
riverine and marine influences affect its presence. Moreover, due to the high human
occupation in the Delta, their influence in terms of distances to hydrologic alteration was
assessed too. The predicted distribution was validated by comparison with a map of actual
habitat type distribution (CORINE land cover) and by field transects. The variables that best
explained the probability occurrence of habitats were soil elevation in habitats with higher
mean elevation (e.g. Cladium-type marshes, dunes/beaches, rice fields and riparian
vegetation) and distance to outer coast in habitats with lower elevations (coastal lagoons, tidal
flats, Salicornia-type marshes and reed beds). The influence of the road and channels on
habitats was reflected in higher soil elevations. The obtained prediction maps have provided
the first results on habitat modelling in the Ebro Delta. The restricted distribution of some
habitats due to human alteration may be the main reason of the mismatch between model
predictions and field data in some habitats.
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1. Introduction
1.1 General context: deltaic system, wetland habitats and human colonization
The Ebro Delta (Catalonia, NE Iberian Peninsula) presents a formidable example of coastal
wetland with a high variability of ecological factors (topography, edaphology, hydrology and
climate) that play a key role in the configuration of its ecological gradients. The confluence of
of contrasted dynamics (i.e riverine, marine and underground) explains most of the Deltaic
variability across spatial and temporal scales. Due to human colonization (settlements,
agriculture, hunting, turism...) wetland habitats with high natural value have been displaced in
the periphery of the Delta where, significant remains of natural ecosystems subsist.
In natural habitats whose sustainability and multi-functional values are threatened, like deltas,
changes in land uses can have more environmental and ecological consequences than in other
ecosystem. Deltaic ecosystems of the Ebro River have particular ecological and economical
value because of their geographic position (interface between terrestrial and coastal zones)
and diversity of habitats (wetlands, coastal lagoons, bays...) (Ibàñez et al. 2010). Human
activities have led to a rapid deterioration of natural aquatic habitats since the beginning of
XX century, mainly, owing to rice cultivation. Until then, however, the dominant landscape
was determined by climatic, geomorphologic and biological factors, except areas transformed
to saltworks. Due to its natural evolution, the Ebro Delta underwent geomorphic changes such
as changes in river mouth and consequent erosion of abandoned lobes, filling of wetlands,
accretion and subsidence of the deltaic plain or regression and coastal progression (Curcó et
al. 1995). These natural processes led rapid modifications of the Delta’s configuration and in
its ecological condition. After several millennia of growth, Ebro Delta was under a river-
dominated dynamics but this trend changed a few decades ago in such a way that the present
Delta is now a wave-dominated coast (Jiménez and Sánchez-Arcilla 1993). This change is
mainly due to the construction of several dams along the river which has caused a nearly total
reduction (97%) of the solid river discharge (Rovira and Ibàñez 2007). Overall, the drastic
reduction of the sediment load slowed down the delta protrusion and intensified the delta
coastline washout (Mikhailova 2003).
7
 
The Ebro Delta contains some of the most important wetland areas in the western
Mediterranean. The majority of the Deltaic plain is devoted to rice agriculture and natural
areas cover only about 25% of the total surface. Among these areas, a set of natural habitats
are present: salt marshes, fresh-brackish marshes, coastal lagoons, sand dunes, natural springs
and bays. Moreover, most of these habitats are included and protected by several European
Directives (e.eg. Habitat Directive and Bird Directive) and regional laws (Ebro Delta Natural
Park). This ecological diversity coexists with a human population near 50.000 inhabitants,
which is located inside the Delta (15.000 inhabitants both Deltebre and Sant Jaume d’Enveja
villages) and along the inner border (approximately 35.000 inhabitants with Amposta and
Sant Carles de la Rapita villages). Urban zones, rice fields and other crops represents near
80% of total Delta surface. Intentisive human colonization in the Delta began at 1860’s with
the first marsh transformations to rice fields after the construction of irrigation channels
(southern hemidelta at 1860 and northern at 1912). From the beginning of 20th century until
present, human transformation of the Ebro Delta largely occurred through draining of
wetlands and the construction of an intensive irrigation system to bring fresh water from the
river to rice fields. According to several authors (Curcó 2006; Mañosa et al. 2001) natural
habitats declined its surface from 27.000 ha (80%) to 11.000 ha (30%) during the 1910-1960
period. The loss of natural habitats stopped during 1960s, but from 1970 to 1990 another 3000
ha of natural habitats were lost, leaving about 25% of total surface still occupied by lagoons
and marshes. Such habitat loss has produced a change in the vegetal and animal communities
present in the Delta. Present management in the Ebro Delta aims at maintaining a high
agricultural productivity and valuable bird populations in the natural areas which are included
in the Natural Park (Ibànez et al. 1997).
1.2 The aquatic habitats of the Ebro Delta
The ecological value of the Delta reflects a high biodiversity, being the aquatic ecosystems
the most important environments which support a representative sample of coastal wetlands.
The Ebro Delta shows 18 natural habitats listed in the annex 1 of the European Directive on
the conservation of natural habitats and of wild fauna and flora (Communities 1991). Bird and
fish populations represent the major faunal groups in order of importance together with
8
 
singularity of halophilous and psammophilous plant communities. There are several technical
reports (Curcó 2006) and scientific studies (Camp and Delgado 1987; Ibàñez et al. 1997;
Menéndez et al. 2002; Valdemoro et al. 2007) that attempt to describe the aquatic
environments of the Ebro Delta for achieving ecological information on their functioning.
These studies note that there is a combination of habitats along a gradient of riverine and
marine influence which confer wide environmental gradients. We consider the next main
habitat units that coexist in the Deltaic plain: estuary, rice fields, coastal lagoons, natural
wells, marshes, dunes and beaches, saltworks, bays and nearshore open sea domain.
Estuaries are dynamic ecosystems that form a transition zone between river environments and
ocean environments. Thus, estuaries are subjected to both marine influences (tides, influx of
saline water...) and river influences (flows, topography of bed...). The pattern of dilution
varies among systems. The Ebro Estuary is a salt-wedge estuary mainly dependent on the
river discharge since the tidal amplitude range is very low (Ibàñez et al. 1997). Ebro Estuary
extends from the mouth to 30 km approximately upstream, the position and presence of the
salt wedge being determined by the tidal range and river discharge.
The rice fields are the dominant landscape of the Delta and all aquatic ecosystems are
influenced by water coming from rice fields. The hydroperiod associated with rice production
is as follows: from April to December, a quantity of ca. 45 m3/s of river water is diverted to
the irrigation canals for continuous irrigation. The resulting eutrophication associated with
large amounts of fertilizer to enhance rice production, as well as pesticides, has led to a
decrease in biological diversity and negative effects on aquatic vegetation of lagoons (Comín
et al. 1991). Nevertheless, the rice fields form an aquatic matrix that link fluvial, lagoon and
marine environments through network channels of irrigation and drainage.
Coastal lagoons are littoral formations formed by the isolation of the marine domain through
the development of a sand bar which separates the water bodies from the open sea (Kjerfve
1994). For their position and origin, their hydrological regime is determined by sea water
inputs coupled with fresh water runoff from drainage of rice fields. For this reason, coastal
lagoons of Ebro Delta show a hydrologic pattern that is clearly reversed (i.e hypersaline
periods do not occur in summer as it would be expected in natural conditions). Aquatic
macrophyte assemblages of the coastal lagoons (e.g. Buda lagoons) have been modified due
9
 
to the creation of salinity gradient that allows the development of different environments with
the coexistence of several species of submerged macrophytes (Comín and Ferrer 1999).
Marshes are wetland areas between terrestrial and marine domains which are linked with the
coastal lagoons. Hydrological settings, mainly water and soil salinity, determine the presence
of fresh (Vilacoto area), brakish (Garxal) and salt marshes (Buda Island) in the Ebro Delta.
Vegetal communities of marshes are adapted to salinity content and soil moisture that
generally depends on frequency and duration of flooding events (Bouma et al. 2005). Marshes
play a key role exhibiting high primary productivity and assist important functions as nutrient
removal and sediment retention (Ibàñez et al. 2002). Under global warming scenario that have
produced eustatic sea level rise of 3.0 – 3,5 mmyr-1 over the past 15 years (IPCC 2001),
scientific literature pointed out the ecological importance of marshes which retain sediments
to offset sea level rise (Day et al. 2000).
The natural freshwater wells are systems of natural ponds situated along the inner Delta
border. In this area, underground water coming from often karstic inland areas (mainly
Montsia) overflows through the surface forming small water bodies no more than 7 metres
deep. A peat layer is present in this zone due to former palustral conditions, being catalogued
as a priority habitat *7210 Calcareous fens in Council Directive 92/43/EEC. These freshwater
wells are popularly called “ullals” and has been considerably altered by human activities,
mainly through draining to lower the underground water level (Capítulo et al. 1994). As a
result, Cladium marshes have been substituted by salt meadows dominated by Juncus genus in
the area of “ullals” de Panxa.
The shoreline of the Ebro Delta is occupied by sandy habitats that contain a very good
representation of dunes and beaches. These habitats also exercise a key role in balancing the
current coastline. The best dune systems of the Delta are placed in la Marquesa and el Fangar.
According to soil mobility, there is a zonation from embryonic dunes (Agropyro-Honckenyion
peploidis), shifting dunes with Ammophila arenaria and fixed dunes (Crucianellion
maritimae). The development of tidal flats with microbial mats in the inner coast of la Banya
peninsula represent another ecological value of these areas with particular hydrological
conditions due to their differential orientation of its coast. Thus, lower topographic levels and
NW dominant winds promote conditions to allow fluctuant moisture along drying and
flooding events.
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The bays are coastal marine waterbodies partially closed with a constant connection to the
sea. In contrast to estuaries, the influence of freshwater is generally limited. In the Ebro Delta,
Fangar and Alfacs bays were originated through the confinement of water bodies due the
formation of spits parallel to the coast. The bays could be considered shallow coastal
ecosystems (2 m. mean depth for Fangar bay and 3,13 m. mean depth for Alfacs bay)(Llebot
et al. 2010), which involve marked spatial and temporal gradients.
The saltworks are traditional salt production areas, located in zones where salt marshes should
have their potential area. This artificial habitat is characterized by deep ponds with variable
salinity that contributes to the increasing diversity of Delta. The Trinitat saltworks were
included in the Natural Park since they provide favorable feeding habitats for the greater
flamingo Phoenicopterus ruber, an emblematic species of the Ebro Delta.
Nearshore open sea habitats are situated along the coastal area in front of the Ebro Delta. This
environment borders on the whole deltaic plain, and the set of beaches, bars and spits are
encompassed by it. A depth of 10 metres is considered the boundary between nearshore and
offshore open sea. Nearshore waters can differ substantially from offshore waters due to the
continental influence, and especially in an estuarine environment like delta. Thus, the latter
are more eutrophic, with higher nutrient and chlorophyll concentrations and different
phytoplankton composition (diatoms and dinoflagellates mainly).
The high diversity of habitats and processes present in the Ebro Delta offers a unique
opportunity to analyse the relationships between Ebro Delta habitats and their environment
and to infer their potential distribution in relation to riverine and marine influences
1.3 Predictive habitat modelling
The term “habitat” has been used in many ways in ecological studies. According to
Spelleberg, (1994) habitat can defined as “the locality or area used by a population of
organisms and the place where they live”. In ecology, the analysis of habitats-environment
relationship has always been a central issue. The major factor involved in habitat distribution,
especially in relation to plant communities, is climate in combination with geology or
hydrology. Habitat factors that are playing a key role in species distribution should be
11
 
considered since species ranges and richness are often correlated with these factors
(Vogiatzakis et al. 2006). The quantification of such relationships represents the core of
predictive geographical modelling in ecology. These models are generally based on various
hypotheses as to how environmental factors control the distribution of species and
communities (Guisan and Zimmermann 2000). Hence, models of habitat distribution are not
subjective models that predict how an area is suitable for development of a particular habitat
in relation to environment conditions.
The relationships between species, communtities or habitats (biotic entities) and
environmental variables are frequently studied using gradient analysis that underlie
hypotheses about species response functions (curves) to environmental gradients (Whittaker
1967). Austin and Smith (1989) defined three types of ecological gradients, namely indirect,
direct and resource gradients. Indirect gradients have no direct physiological influence on
species performance (slope, aspect, elevation, topographic position, geology). Direct gradients
are environment parameters that have physiological importance, but are not consumed (e.g
temperature, pH). Resource gradients address matter and energy consumed by plants or
animals (nutrients, water, light, food for plants, water for animals…). Generally, literature
pointed out that indirect variables usually replace a combination of different resources and
direct gradients in a sample way (Guisan et al. 1998; Guisan et al. 1999).
The real or actual vegetation is a patchwork of different classes or categories of communities.
Classifying these communities in accordance with some key allows one to construct a
vegetation map, which can be interpreted of actual vegetation that is normally more complex
than habitat units map. While they are a simplification of reality, habitat maps are important
data for correct environment management of a territory. The Ebro Delta has an important
environmental dataset of which habitat maps are included. This work derived from the
adaptation of the CORINE Biotopes project in Catalonia (Carreras and Diego 2007). But, this
kind of information (i.e habitat maps or vegetation maps) has a limited temporal variability.
Even in absence of human influence, vegetation dynamics is complex and intense, especially
in deltas which are subject to sharp environment gradients. In the Ebro Delta, the ecological
term of succession is applicable, defining the natural sequence in which a habitat replaces
another over the passage of time. Then, the potential vegetation is defined as the stable
community which would exist in an area as a consequence of progressive geobotanical
succession if man ceased to affect and alter the terrain. Curcó et al. (1995) made an exercise
12
 
to delineate potential vegetal domains based on topographical and sedimentological features
of the Deltaic plain. In this case, potential habitats will be more in balance with the salinity
and moisture conditions of the environment. A model that considers sites with their
disturbance features (e.g road or channels) might be expected to explain only a portion of the
variance in habitat type distribution. Even so, this approach can be a chance for applying in
the Delta.
The first step that has to be considered in predictive modeling is the link between habitat units
and mapped physical data. Several modelling methods have been used in scientific papers:
heuristic, decision trees and statistical methods. The last approach, mainly regression, is the
one most used to predict the value of the response variable if continuous, or the probability of
a variable if categorical (Vogiatzakis 2003). Most predictive modelling efforts has used
logistic regression to predict species (Rüger et al. 2005), vegetation assemblages (Davis and
Goetz 1990) or animal habitat (Corsi et al. 1999). A logistic regression is well-suited where
the dependent variable is dichotomous (presence/absence of habitats), and the technique
allows one predictor (binary logistic regression) or more than one (multiple logistic
regression). In addition the method lets a non-Gaussian distribution of the independent
variables (Hosmer and Lemeshow 2000). Also, the result of the regressions ranges from 0 to 1
so that is appropriate for the generation of a likelihood model (Álvarez-Arbesú and Felicísimo
2002). The application of this method to wetlands and aquatic ecosystems is not an exception.
(Narumalani et al. 1997) applied multiple logistic regression to predict aquatic macrophyte
distribution. Another similar study was applied to aquatic vegetation by van de Rijt (1996) for
predicting vegetation zonation in a former tidal area, while Shoutis et al. (2010) applied it for
predicting riparian vegetation based on terrain variables and different river orders. The final
step to consider in a predictive model is the model validation. Such evaluation consists in
determining the suitability of a model for specific applications. According to Pearce and
Ferrer (2000), wherever possible, evaluation is best undertaken with independent data
collected form sites other than those used to develop the model. If independent data are not
available, there are statistical techniques that fit the model in different degree, such as receiver
operating characteristic (ROC) plot methodology (for more details see methods section). The
next figure schematizes the generic steps of predictive modelling (Figure 1):
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Fig. 1. Generic steps of predictive habitat modelling.
The most used environmental predictors in predictive habitat modelling studies are those
related with topography, geology and climate (Franklin 1995). Topography and its attributes
such as elevation and slope, derived from Digital Elevation Model (DEM) are among the
principal variables employed in these works due to their importance on vegetation patterns. A
digital elevation model is any digital representation of the continuous variation of relief across
space (Burrough et al. 1998). The use of accurate DEMs is especially important for deltas
because, in the case of the Ebro Delta, about 40% of the plain surface lies under 0,5 meters
above mean sea level (Ibàñez et al. 1996). In addition, hydrological variables such as
frequency and duration of inundation are the main limiting factors of lagoon-wetland
complex, and can be inferred through differences in soil elevation (Hickey and Bruce 2010).
It is important to note that DEMs frequently contain systematic errors which can limit the
effectiveness of predictive habitat distribution. On the other hand, too high accuracy will
detect microtopography relief that may lead to unsatisfactory results.
Ecological data sets have two distinct characteristics when compared to other kinds of data:
they are multivariate and location specific. Recent studies to predictive modelling of habitats
have been developed on a Geographic Information System (GIS), and ecological modellers
have focus on incorporating spatial patterns in the models to apply them in large geographic
areas (Vogiatzakis and Griffiths 2006; Zare Chahouki et al. 2010). Ecological modelling with
GIS involves its complementary use for addressing ecological approaches, such potential
distributions. In scientific literature, there are two ways of linking ecological models with
GIS: 1) run the model outside the GIS and use the GIS for pre-processing data set (e.g.
Statistical analysis Predictive modelling
Model validation
Environment
Habitats
14
 
coordinate system transformation, location of sample points…) and generate cartographic
outputs and 2) use GIS for extract metrics on environment mapped variables which will
conforms the core of statistical method and post-processing of the data through cartographic
display too (Felicísimo 2003; Felicísimo et al. 2002; Franklin et al. 2000). GIS-based spatial
analysis tools facilitate the representation of ecological data across the space and its
correlation with environmental data. In deltaic environment (among marshes, lagoons...),
surface elevation (or water depth) and inundation frequency are the most important
environment variables for vegetal zonation (Silvestri et al. 2005). Spatial analysis provides
tools for researchers to assess how these factors influence habitat type distribution, extract
metrics and explore the relationship between aquatic environments and topography by
investigating species zonation. For example, Hickey et al. (2010) examined the relationship
between distribution of salt marsh vegetation and the extent of tidal inundation using fine
elevation data; (Xie et al. 2011) defined several landscape units of freshwater wetlands in
Florida based on surface elevation; (Moran et al. 2008) linked spatial variation of flooding
regime with the vegetation zones in a karst wetland.
Habitats of the Ebro Delta are along environmental gradients which can be assessed to infer
distribution patterns. Getting a predictive model, one can establish a relation between the
habitat units and the environment data. The geographic scale of the Delta offers the
opportunity to incorporate Geographic Information System techniques for extrapolating over
a wide range those relations.
 
 
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2. Hypotheses and objectives
Our initial hypotheses are: i) habitats of deltas are distributed spatially as a consequence of
specific environmental requirements, mostly surface elevation and distance to sea/river. ii)
These relationships (between delta habitats and surface elevation and the distance to sea/river)
can be used to build a model that describes the potential distribution of each habitat according
to the present configuration of deltaic plain.
The main objective of this study was to determine the potential distribution of some existing
wetland habitats in the Ebro Delta through a predictive habitat model based on terrain
variables. To achieve this aim, the following specific objectives have been proposed:
- To get elevation ranges of each habitat type within the altitude gradient of the delta
by a digital elevation model (DEM). To validate them with field data.
- To calculate distance ranges from the geographical position of each habitat type
relative to the river and marine influence, which are determined from delta hydrologic
boundaries.
- To apply the predictive model in a Geographic Information System (GIS) to obtain
maps of probability of presence for each habitat.
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Research questions
By achieving the objectives of the exercise, one would be able to answer the following
questions:
- What are the most important environment variables that determine the potential
distribution of the main habitats in the Ebro Delta?
- Are the human factors more important predictors than distances to hydrological
boundaries in explaining the distribution patterns of wetland habitats? Or conversely, are the
terrain variables?
- The spatial distribution of aquatic habitats could be predicted accurately by
developed predictive model?
- Is the statistical technique of logistic regression appropriate to predict the link
between physical variables and aquatic habitat types?
- What are the ranges of elevation and distance to the hydrological boundaries of the
delta of each habitat type?
Research strategy
This work is part of a broader study of palaeocological reconstruction of the Ebro Delta based
on geochemical and biological analysis of its sediments. Two major steps can be
distinguished within the research strategy of this Master Thesis: (1) modelling the link
between the biological data and the accompanying terrain physical data and (2)
implementation of the model in a Geographic Information System environment in order to
attain a coverage habitat predictive map in terms of probability of occurrence for each aquatic
habitat. These early steps will provide basic ecological information for evaluating the
relationship between the types of aquatic habitats and the biological proxies (mainly fossil
diatoms) preserved in its sediments.
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3. Methods
3.1 Study area
The study was carried out in the Ebro Delta, which is one of the largest deltas in the
northwestern Mediterranean, with 330 km2
(Figure 2). Within this extension, rice fields
occupy the majority of the delta plain (65% of the total surface) and natural areas cover only
about 80 km2
(25%). These areas include a variety of aquatic habitats, providing an excellent
example of coastal wetlands habitats: riparian vegetation, salt, brackish and fresh water
marshes, coastal lagoons, natural springs, bays, sand dunes and mudflats. The confluence of
contrasted dynamics, mainly riverine and marine, explains most of this high spatial
variability. This diversity provides the presence of a large number of habitats of community
interest listed in the annex 1 of the European Directive (Communities 2003) on the
conservation of natural habitats and of wild fauna and flora. The best preserved natural areas
are included in the Natural Park of the Ebro Delta that comprises 7.802 ha. Other zones that
also include part of the rice fields are protected under other regulations of the Catalonian
Government and the European Union (i.e Natura 2000).
18
 
 
Fig. 2. Location of the study area, the Ebro Delta.
 
There were various reasons for choosing this study area. Firstly, there is good information
available, both biological (habitat maps) and physical data (digital elevation model).
Secondly, there is a great spatial heterogeneity, favouring the existence of diverse
environments, and allowing a wide range of ecological gradients to be assessed with the
actual deltaic plain configuration. And thirdly, there is a wide set of aerial and topographic
maps with different scales for carrying the GIS dataset analysis.
The Ebro Delta shows a very low relief, with slopes of around 0,01 – 0,02 %. Riverine and
sedimentary dynamics determine a elevation gradient decreasing from lévées in the inner
border (4-4,5 m) to the river mouth (0-0,5 m) (Figure 3). Lévées of former river arms have
more elevation than the adjacent deltaic plain and these structures can be recognised in the
topographical maps. The low elevation areas are usually the ones having more marine
influence. The agricultural activity has been the major factor modifying of the native
19
 
topography of the delta, either lowering high areas or filling lagoons. Moreover, currently the
Ebro Delta is undergoing elevation loss due to sediment deficit created by sediment retention
in the dam system along the river. This fact tends to lead a negative balance between vertical
accretion and subsidence on in the deltaic plain. Furthermore, elevation loss is accelerated by
sea level rise due to the effects of climate change.
300000
,000000
300000
,000000
320000
,000000
320000
,000000
4500000
,000000
4500000
,000000
Elevation (m)
0 - 0,5
0,5 - 1,0
1,0 - 1,5
1,5 - 2,0
2,0 - 2,5
2,5 - 3,0
3,0 - 3,5
3,5 - 4,0
> 4,00 5 102,5
Km.
´
 
Fig. 3. Digital Elevation Model of the Ebro Delta. Source: Cartographic Institute of Catalonia, 2011.
3.2 Wetland habitats, terrain variables and hydrologic alterations
The natural habitat classification and mapping of the CORINE Biotopes (Communities 1991)
developed for Catalonia was used to identify and select each wetland cover type. This data
source includes two types of information, 1) the list of CORINE habitats of Catalonia (Vigo et
al. 2006) and 2) the mapping of habitats in Catalonia 1:50,000 (Carreras and Diego 2007).
The list has a hierarchical structure based on habitats classification of annex 1 of European
Union Habitats Directive and describes each habitat unit from physiognomical, ecological and
phytosociological characters. Overall, 9 habitat types have been selected to develop the
model. The selection of wetland habitats responds to different criteria as a function of the
20
 
variability on hydrological requirements and salinity tolerances. Since the Ebro delta is a
coastal system, the distribution patterns of broad types of wetland habitats, such as
Salicornia-type marshes or salt meadows can be influenced by salinity. Without freshwater
inputs, topography should be the main factor determining the habitat distribution. It is known
that flooding regime is a primary factor structuring coastal wetlands with the frequency and
duration of inundation determined by surface elevation (Hickey and Bruce 2010). In addition,
the geographical position of each habitat respect fluvial and marine influence will affect its
distribution in the deltaic landscape. In this study we have assessed how the target habitats are
distributed through a combination of several distances that are related with the hydrologic
boundaries of the delta plain (see section 3.5). However, the effects of the hydrologic
alterations produced by two main anthropogenic sources should also be taken into account,
these being 1) fresh water inputs due to irrigation from adjacent rice fields and the network
irrigation channels; 2) roads that can interfere natural hydrologic fluxes. Thus, distances to
these hydrologic alteration sources have been included as well in the model as possible
predictors of geographic distribution of the wetlands habitats.
3.3 Dependent variable: current distribution of wetland habitats
The presence or absence of the wetlands habitats has constituted the dependent variable of the
model. For this purpose, the map of natural habitats on 1:50.000 scale has been used. This
data set was acquired through digital format (shape file on ArcView environment) from the
Environment Department of the Government of Catalonia. The sheets that cover the Ebro
Delta include numerous habitats, from the dune domain to reed beds; at the same time each
polygon comprises several classifications. In this study we chosed the main wetland habitats
present in the Ebro Delta and the final delimitation of target habitats was subject to expert
review. Most of the habitats (7 out 9, except reed beds and rice fields) are classified as
community interest by the European Union Habitats Directive. The directive defines habitats
of Interest as those that (i) are in danger of disappearance in their natural range; or (ii) have a
small natural range following their regression or by reason of their intrinsically restricted
area; or (iii) present outstanding examples of typical characteristics of one or more of the
21
 
seven following biogeographical regions: Alpine, Atlantic, Boreal, Continental,
Macaronesian, Mediterranean and Pannonian.
The Interpretation manual of European habitats (Romao 1996) was used to describe each
wetland habitat from digital maps of the Ebro Delta. The list in table 1 shows the habitats
included in the study and its corresponding classification according to CORINE Biotope
classification, and it also lists the most representative sites of Delta where these habitats are
present. So, this classification system has resulted in the list of habitats of Catalonia. The
codification system of habitats is based on a hierarchical classification and has been identified
by a code like nn.xxxx, where the first two digits indicate the main group it belongs to (Table
2). Thus, the code of each habitat provides information on the groups and subgroups to which
they belong and with which other habitats have similarities.
Table 1. Main CORINE groups of European habitats classification.
Habitat CORINE group
Coastal and halophytic communities 10
Non-marine waters 20
Shrubby vegetation and grassland 30
Forests 40
Bogs and marshes 50
Screes 60
Agricultural land and artificial landscapes 80
Burned areas 90
22
 
Table 2. Wetland habitats included in the study and its corresponding classification based on CORINE
biotope project. * Priority habitat.
Wetland habitat Code HCI CORINE code Delta sites
1. Coastal lagoons *1150 Coastal lagoons 21 Lagoons Encanyissada,
Tancada,
Aufacada,
Platjola, Illa de
Buda, Garxal,
Canal Vell, les
Olles
2. Sandy habitats 2110 Embryonic shifting
dunes
2120 Shifting dunes along
the shoreline with
Ammophila arenaria
(white dunes)
2210 Crucianellion
maritimae fixed beach
dunes
16.1 Sand beaches
16.2 Dunes
Along shoreline
of the Delta plain
3. Tidal flats 1140 Mudflats and
sandflats not covered by
seawater at low tide
14 Mud flats and sand
flats
La Banya, Fangar
4. Salicornia-type
marshes
1420 Mediterranean and
thermo-Atlantic
halophilous scrubs
(Sarcocornetea fruticosi)
15.6 Halophilous
shrubby formations
Buda island
(Calaixos),
Tancada
5. Salt meadows 1410 Mediterranean salt
meadows (Juncetalia
maritimi)
15.5 Mediterranean salt
meadows
Sant Antoni,
Garxal, Tancada,
Encanyissada
6. Cladium-type marshes *7210 Calcareous fens
with Cladium mariscus
53.3 Cladium mariscus-
dominated formations
Vilacoto, Ullals of
Baltassar
7. Reed beds - 53.1 Reed beds Garxal,
Encanyissada,
Tancada, Platjola,
Aufacada, Canal
Vell, les Olles
8. Rice fields - 82d Rice fields Over the deltaic
plain except
peripheral areas
9. Riparian vegetation 92A0 Salix alba and
Populus alba galleries
44.1 Riparian willow
formations
Sapinya island
23
 
Legend
Coastal lagoons (1150)
Sandy habitats (dunes and beaches) (2110, 2120, 2210)
Tidal flats (1140)
Salicornia-type marshes (1420)
Salt meadows (1410)
Cladium fens (7210)
Reed beds (53.1)
Rice fields (82d)
Riparian vegetation (92A0)
Ebro river
Human settlements0 5 102,5
km.
´
Fig. 4. Map of
habitats of the Ebro
Delta with its
corresponding
CORINE code
24
 
Description of wetland habitats
1. Coastal lagoons
EU habitat code: *1150; CORINE code: 21
Coastal lagoons of the Ebro Delta are typical water bodies from a deltaic environment
(albufera-type) formed due to the evolution of the Delta lobes. Because of their origin
and separation from the open sea by a sand bar, they are lagoons strongly influenced by
seawater inflows. In their original state they were salt water lagoons with a maximum
salinity in summer, but due to the rice field drainage, their hydrologic regime has been
severely. Coastal lagoons are zones with high biological and ornithological importance,
where several species listed in the Bird European Directive are present. The aquatic
vegetation of the coastal lagoons is composed of mixed macrophyte beds of Ruppia
cirrhosa, Potamogeton pectinatus and Zostera sp. (Menéndez et al. 2002). There are a
total of nine coastal lagoons in the delta, among them Buda island, placed near the river
mouth, or the Aufacada lagoon.
 
Fig. 5. Present distribution of the coastal lagoons in the Ebro Delta according to the habitat
mapping of Catalonia. Source: Department of Sustainability and Territory, Catalonia
Government.
25
 
2. Sandy habitats (dunes and beaches)
EU habitat code: 2110, 2120, 2210; CORINE code: 16.1, 16.2
These habitats are basically constituted by deltaic-front sand bodies. Ecologically sandy
habitats can be characterized as environment that contents low water content, low levels
of salts and organic matter and relative levels of mobility. Sandy habitats of the Ebro
Delta bring together three types of habitats of community interest related to the
substrate mobility: embryonic dunes (2110), shifting dunes with Ammophila arenaria
(2120) and fixed dunes (2210). These habitat types are considerate as transitional and
littoral sedimentary environments due to marine agents produce largely the mobilization
of its soils. Then, they are associated with littoral transfer process. The extension of this
habitat in both hemideltas is unequal. The main reason is the different orientation of the
outer coast with respect to prevailing winds (NW). Thus, the most representative area of
beaches and dunes systems is located in the northern hemidelta: Marquesa beach-Garxal
and Punta del Fangar.
 
Fig. 6. Present distribution of the sandy habitats (dunes and beaches) in the Ebro Delta
according to the habitat mapping of Catalonia. Source: Department of Sustainability and
Territory, Catalonia Government.
26
 
3. Tidal flats 
EU habitat code: 1140; CORINE code: 14
Flat coastal areas, devoid of terrestrial vascular plants and usually colonised by blue-
green algae and diatoms. This habitat occupies coastal sands and muds and their
associated coastal lagoons that experience recurrent episodes of flooding and drying. It
is particularly well developed and forms the greatest extension in the Alfacs Peninsula,
formed by la Banya spit and Trabucador barrier. This area is very sandy, and flooding
periods are frequent due the strong northwestern winds, which results in a vertical
stratification of physicochemical gradients between the aqueous interface and the solid
substrate (Mir et al. 2000).
 
Fig. 7. Present distribution of the tidal flats in the Ebro Delta according to the habitat mapping
of Catalonia. Source: Department of Sustainability and Territory, Catalonia Government.
27
 
4. Salicornia-type marshes
EU Habitat code: 1420; CORINE code: 15.6
Low shrubby expanses of woody glassworts which in the Ebro Delta are dominated by
succulent perennial species of the genus Sarcocornia and Arthrocnemum. Within the
water salinity gradient of marshes, salt marshes are the wetlands with major influence of
marine water. In them, the connexion to freshwater is limited, except for those zones
that are receiving water inflows from of adjacent rice fields. Depending on rainfall,
evaporation and tidal exchange, the salinity pattern may differ through the year. The
differences of these factors can influence the ecological and physical traits of each
marsh, such us vegetal communities (halophytic and hydrophytic), net primary
productivity or accretion and subsidence rates. Buda Island is the most representative
Arthrocnemum-type marsh in the Delta.
 
Fig. 8. Present distribution of the Salicornia-type marshes in the Ebro Delta according to the
habitat mapping of Catalonia. Source: Department of Sustainability and Territory, Catalonia
Government.
28
 
5. Salt meadows
EU Habitat code: 1410; CORINE code: 15.5
This habitat is characterized by the presence of Juncus acutus and Juncus maritimus as
the most representative plant. These taxa withstand high soil humidity and for this
reason grow in drenched and/or periodically submersed soils. However, the habitat finds
its ecological optimum in sites occurring at least a few centimetres higher than the
average soil water level. In the Ebro Delta, it grows in scattered inland sites where soil
elevation is higher than those of the halophilous scrub. Regarding salinity, this habitat
forms a transitional stage between salt marshes Salicornia-type and habitats lacking
halophytic vegetation. In the Ebro Delta, the salt meadows can form intermediate stands
with halophilous scrubs. According to Curcó et al. (1995) this terrestrial habitat have
been drastically reduced in relation to their potential surface area since that area has
been impounded by the rice fields.
 
Fig. 9. Present distribution of the salt meadows in the Ebro Delta according to the habitat
mapping of Catalonia. Source: Department of Sustainability and Territory, Catalonia
Government.
29
 
6. Cladium-type marshes
EU Habitat code: 7210; CORINE code: 16.2112
This habitat type is the only one the wetland habitats considered that constitute a
priority habitat in the Ebro Delta. Within the fresh water ecosystems, the presence of
Cladium-type marshes was originally associated with underground freshwater springs in
karstic zones (Ullals) or in elevated zones with recurrent flooding events. Nowadays the
most representative zone of this habitat in the Delta is in the Vilacoto area at the east of
the Encanyissada lagoon. In this habitat the presence of dense helophytic communities
dominated by Cladium mariscus, Phragmites australis and Scirpus maritimus is linked
with a superficial peat layer and a significant input of underground water; this fact allow
the submersion of the base of the plant during most of the year.
 
Fig. 10. Present distribution of the Cladium-type marshes in the Ebro Delta according to the
habitat mapping of Catalonia. Source: Department of Sustainability and Territory, Catalonia
Government.
30
 
7. Reed beds
EU Habitat code: - ;CORINE code: 53.1
The habitat occurs near lagoons, channels or other wetland types which receive direct
fresh water influence from the rice fields and the river. It occurs in still, fresh or
brackish water. Within the Ebro Delta, natural colonies of Phragmites australis develop
in the Garxal area, which is subjected to the direct influence of the riverine processes.
Along the south edge of the lagoon there is an intermediate belt of brackish reedswamp
dominated by Phragmites and Juncus species. Over the Delta plain, this habitat has
spread along the margins of the coastal lagoons and bays due to hydrological changes
caused by rice cultivation mainly.
 
Fig. 11. Present distribution of the reed beds in the Ebro Delta according to the habitat mapping
of Catalonia. Source: Department of Sustainability and Territory, Catalonia Government.
31
 
8. Rice fields
EU Habitat code:- ; CORINE code: 82d
This habitat is the dominant landscape of the Delta as a result of a large agricultural
occupation process that has led an actual coverage of near the 70% of the deltaic plain.
Despite being a humanized environment and classified as artificial landscape for
CORINE Biotope project, the rice fields constitute a aquatic matrix that link fluvial,
lagoon and marine ecosystems. During the rice inundation period (May-December) this
habitat acts as an authentic aquatic ecosystem which offers zones of feeding and resting
to aquatic birds. Nevertheless, the inflow of huge amounts of fresh water into the fields
has been an important factor in alterating the hydrology of the Delta, as well as causing
loss of wetlands habitats and loss of elevation of the deltaic plain.
 
Fig. 12. Present distribution of the rice fields in the Ebro Delta according to the habitat mapping
of Catalonia. Source: Department of Sustainability and Territory, Catalonia Government.
32
 
9. Riparian vegetation
EU habitat code: 92A0; CORINE code: 44.1
This type of habitat is found growing in the Sapinya Island. According to CORINE land
cover, this is the only patch of riparian vegetation present in the Ebro Delta. Under
natural conditions, the lower Ebro River was bordered by riparian forest along its
levees. Generally, this habitat types inhabited in the fluvial levees in mean elevation
range of 2 and 4 m above sea water level. Flooding events in these areas only occurred
when the river overflowed large flows, but due to construction of the dam system along
Ebro river watershed the river flow has been drastically laminated. Coupled with human
colonization of delta plain in terms of agricultural purposes, which was more significant
in these higher zones, the riparian habitats have a relictual distribution.
 
Fig. 13. Present distribution of the riparian vegeation in the Ebro Delta according to the habitat
mapping of Catalonia. Source: Department of Sustainability and Territory, Catalonia
Government.
33
 
3.4 The independent variables: elevation and distances to hydrologic boundaries
The Ebro Delta and the Mediterranean deltaic systems generally have a complex
structure and its functioning depends on hydrologic, geologic and climatic factors. The
diversity of habitats in the area of study is high, forming a set of environments that are
river- and marine-dominated. The first factor has lost importance due to the dam
construction along the Ebro River watershed concerning the reduction of near the 99%
in the particulate sediments of the lower river. Today, the hydrological conditions in
some salt marshes of the Delta are dominated by inputs of seawater through outlet
channels, much more than riverine influence, being the agricultural runoff the factor that
is altering the natural conditions (except in the river mouth area, Garxal). The
topographical factor plays a key role in the Ebro Delta since about 40% of the plain
surface lies under 0,5 meters above mean sea level (Ibàñez et al. 1996). In addition,
hydrological factors are highly correlated with the variation of soil elevation that will
determine frequency and duration of the inundation events. The distribution and
composition of lagoon-marshes complexes depends strongly on this terrain variable.
Thus, river lévées are the highest parts of the Delta, and under natural conditions, are
vegetated by riparian forests such as Populus and Salix galleries. These habitats are
flooded only during high discharge. Outside these areas are fresh, brackish or salt
marshes, depending on factors such elevation, inputs of upland runoff, riverine
influence, marine influence or soil drainage. Regarding vegetation marsh zonation, in
some cases there is a clear vegetation transition related with soil salinity and water
regime (Silvestri et al. 2005). The link between these terrain factors and vegetal
communities is one of the research questions of this study.
The independent variables included in this study for assessing the potential distribution
of the wetland habitats have been surface elevation, distance to hydrologic alterations
and distance to river/sea influence. The last approximation was assessed by the
combination of several distances which are associated with the hydrologic boundaries of
the Delta plain and will serve to extract influence of the flooding regime as an indirect
way. The hydrologic alteration approach includes all of the elements on the deltaic
34
 
landscape that have resulted from human activity, mainly roads, irrigation channels and
rice fields.
Table 3. List of terrain predictors included in the study.
Terrain variable Abbreviation Description Source
Surface elevation ALT Terrain altitude of the delta
plain
Digital elevation model 1x1 m
(Base cartogràfica de l’Institut
Cartogràfic de Catalunya)
Distances to
river/sea
influence
- Outer coast OC External shoreline Topographical map
1, 25.000 (ICC)
- Inner border IB Inner side of deltaic plain Topographical map and
ortophotomaps (ICC)
1, 25.000
- River channel RC Ebro river course and its
levees
Topographical map
1, 25.000 (ICC)
- Lagoon LAG Coastal lagoons Topographical map
1, 25.000 (ICC)
- Bay BAY Coastal marine water bodies
partially closed
Topographical map
1, 25.000 (ICC)
- Former river
arms
FR Ancient river courses: riet
Fondo, riet de Zaida and riet
Vell
Digital elevation model
(ICC)
- River mouth RM Current mouth of the Ebro
river
Topographical map
1, 25.000 (ICC)
Distances to
hydrologic
alteration
- Roads ROAD Roads and paths constructed
over the Delta plain
Topographical map
1, 25.000 (ICC)
- Channels CHAN Network of channels for
irrigation and drainage
waters from rice fields
Topographical map
1, 25.000 (ICC)
- Rice fields RF Agricultural crops of rice CORINE land cover (Department of
Sustainability and Territory)
35
 
Legend
Ebro river
Coastal lagoons
Bay
Former river arms
River mouth
Outer coast
Inner border0 5 102,5
km.
´
Fig. 14. Map of
hydrologic boundaries
from river/sea
influences in the Ebro
delta.
36
 
Legend
Ebro river
Coastal lagoons
Channels
Roads
Rice fields5 0 52,5
km.
´
Fig. 15. Map of
hydrologic alterations
elements from human
colonization in the Ebro
delta.
37
 
3.5 Vegetation transects
In order to validate the soil elevation of wetland habitats obtained via CORINE land
cover, transects that cover soil elevation gradient has been developed by mean transects.
Along transects, the presence of each habitat through the recognition of homogenous
belts was recorded and sample points were georeferenced (European Datum 1950, UTM
31N). The next habitats were surveyed: Salicornia marshes, Juncus marshes and reed
beds (fresh-brackish marsh) (Figure 16). Transects of salt marshes of Salicornia-type
were developed in Sant Antoni Island, a marine-influenced area of Buda Island. In this
site, succulent Salicornia and Juncus often co-occur. The Garxal brackish marsh
bordered lagoon which receive river discharge directly. Transects were develop along
south edge of the lagoon, where a belt of Phragmites marshes is present. The habitat of
salt meadows dominated by Juncus genera was located in la Tancada area and la
Platjola. Salt meadows of la Tancada are the area of Delta coincident with CORINE
habitat map since in other marshes it were been detected (i.e. Garxal marsh) but it didn’t
incorporated in the digital maps.
Ebro river
Coastal lagoons5 0 52,5
km.
´
Tancada: Juncus and Salicornia marshes
Sant Antoni
island: Salicornia
and Juncus
marshes
Garxal marsh: Phragmites and
Juncus marshes
 
Fig. 16. Location of study of marsh study sites in the Ebro Delta where elevation transects were
developed.
38
 
3.6 GIS development
Geographic Information Systems (GIS) are widely used for ecological studies because
they provide techniques to relate several environmental/landscape variables and their
specific location. This approach allows to assess the distribution of habitats according to
environmental gradients and to suggest correlations between them. The first step has
consisted to obtain the database that integrates the main variables included in the study:
current distribution of wetland habitats and terrain deltaic predictors. Therefore the
process was sequential: initially a database was established and then the model was
applied for establishing the relationship between the wetland type distribution and
terrain variables. A flow diagram of the general process is presented in Figure 17, and
the specific cartographic methods will be explained in the following sections.
Habitat maps
Coastal lagoons, pres/abs
Topographic maps
Ortophotomaps
Digital elevation model
Distances to river/sea
influence
Distances to
hydrologic alteration
Terrain predictors
Distribution of wetland habitats
Tidal flats, pres/abs
Wetland type, pres/abs
.
.
.
Logistic regression
Predictive wetland
habitat model
Random sampling points,
pres/abs
Predictive habitat
mapping
Validation
 
Fig. 17. Flowchart of the process of generating the predictive model (grey boxes) and the
current distribution of wetland habitat types of the Ebro Delta.
39
 
Database development
Wetland habitat data were collected following a selection of habitat type layers using
the CORINE land cover as vector format. A Geographical Information System (GIS)
has been developed to generate the database of wetland habitat types (presence and
absence) in the Ebro Delta. For this purpose, the commercial GIS package ARC/INFO,
ArcView 9.3 was used. The cartographic data sources required to generate wetland
covers and terrain variables are listed in the follow table (Table 4). All of the layers
have been projected in UTM north zone 31, European Datum 50.
Table 4. Data sources for obtaining digital database of wetland types and terrain variables.
 
Layer Format Scale/
Precision
Output layers Source
CORINE land
cover
Polygon,
Shapefile
(ArcView)
1, 50.000 - Pres/abs of
each habitat
type (n = 9)
- Rice fields
Department of Sustainability
and territory (Government of
Catalonia)
Digital Elevation
Model
ASCII, raster
(ArcView)
1x1 m - Raster
elevation grid
- Former river
arms
Cartographic Catalan Institute
(ICC)
Topographical
map
Polygon,
Shapefile
(ArcView)
1, 25.000 - Ebro river
- Lagoons
- Bays
Cartographic Catalan Institute
(ICC)
Polygon,
shapefile
1, 25.000 - Buildings Cartographic Catalan Institute
(ICC)
Polyline,
shapefile
1, 25.000 - Outer coast
- Irrigation
channels
- Roads
Cartographic Catalan Institute
(ICC)
Ortophotomaps MrSID, raster 1, 5.000 - Deltaic plain
- Inner border
- River mouth
Cartographic Catalan Institute
(ICC)
40
 
The complete cartographic procedure for obtaining the database is shown in Figure 18,
but the general steps can be summarized as follow:
a. Digitalization of the Delta plain to obtain the geographical limit of the habitat
and distance layers.
b. Obtaining the habitat map cover for each target wetland (vectorial format):
coastal lagoons, salt meadows, Salicornia-type marshes, Cladium marshes, Reed
beds and rice fields. The habitat map cover of sandy habitats (dunes and
beaches) and tidal flats were modified from the original sheets through expert
criteria.
c. Extraction from the topographic sheets (1,25.000 scale) the hydrologic layer in
vectorial format (polygon and polyline). From the polygon data, select by
attributes the cases of river courses (RC), lagoons (LAG) and marine waters
(BAY). From the polyline data, select by attributes the cases of outer coast (OC)
and channels (CHANN).
d. Extraction from the topographic sheets (1,25.000 scale) the settlement layer in
vectorial format (polygon and polyline). From the polygon data, select by
attributes the cases of generic buildings (BUIL). From the polyline data, extract
by attributes the cases of roads (ROAD).
e. Digitalization of the inner border (IB) and river mouth (RM) in polyline format
from the ortophotomaps on 1, 5.000 scale.
f. Recognition and digitalization of the former river arms (FR) from the DEM of
Delta.
g. From each wetland habitat map polygons with habitat presence, elevation
metrics (min, max, range and mean) were extracted with zonal statistics tool of
the Analysis tools extension.
h. From each wetland habitat map polygons with habitat presence, distances to
closest feature of hydrologic and anthropogenic layers was calculated with near
tool of the Analysis tools extension.
41
 
Ortophotomaps
Habitat maps cover
Raster data Vector data
Inner border (IB)
Digit.
Digit.
Topographic maps Hydrologic polygon layer River channel (RC)
Channel (CHAN)
Outer coast (OC)
Lagoons (LAG)
Bays (BAY)
Select by attributes
CAS = MAI003
CAS = MAI001
CAS = MAI011
Hydrologic polyline layer
Select by attributes
CAS = CAN001
Clip
Clip
CAS = CNA
Settlement polyline layer Roads (ROAD)
Select by attributes
CAS = VIA
Clip
DEM
Digit.
Former river arms (FR)
Zonal statistics as a table
Elevation metrics tables
Elevation metrics tables
Elevation metrics tables
Elevation metrics tables
Elevation metrics tables
…
Delta
River mouth (RM)
Clip
Near
Coastal lagoon pres/abs map
Tidal flat pres/abs map
Salicornia-type pres/abs
mapsSalt meadows pres/abs maps
…Rice fields pres/abs maps
RC
LAG
BAY
IB
OC
FR
RM
ROAD
CHANN
RICE
Distance to terrain
variables tables
Settlement polygon layer
CAS = EDI001
Buildings (BUI)
Clip
Fig. 18. Cartographic
model to assess the
current distribution
of wetland habitats
42
 
Generating data for the predictive model
Once the dataset of habitat maps was developed, the next step was to create a random
sample of 1.000 points for each wetland habitat type over the whole Delta plain to
achieve data of the presence/absence of the habitats. The process was sequential again,
and the steps are specified in the cartographic model (Figure 19). Several layers
obtained in the previous steps (see database development) have been taken into
consideration as excluding layers to overlap with the sample points. These geographic
areas are: river channel, lagoons, irrigation channels, roads, buildings and rice fields.
We excluded theses areas while the model was developed because they are parts of the
delta where the habitats will be neither present and absent due to its land use. To sum up
the process, the following steps are listed:
a. Merging shapefiles of infrastructure elements (roads, channels and generic
buildings) to obtain the constraining layer.
b. Merging the infrastructure layer with hydrological layers (river channel, lagoons
and rice fields).
c. Creating a random point dataset (n = 1000) along the Delta as geographical limit
layer. For each habitat type a set of layer random points was created (n = 9).
d. Overlap data point layer with the excluding layer (infrastructures + water). The
output layer was defined as points where the habitat is not present (layer absence
points).
e. For obtaining the layer of presence points of each habitat, a new random point
dataset with the same number of absence points (to avoid bias in the sampling
process) was created. The limit layer was the specific habitat presence map.
f. Rasterization of vector layers for obtaining digital data of the independent
variable over whole Deltaic plain: distances to rive/sea influence and
anthropogenic limits. For this purpose the Euclidean distance tool of Spatial
analysis extension was applied for each layer with 1m of pixel resolution.
g. Each sample point layer (presence and absence) has its respective value of
elevation, distances to river/sea influence and distances to hydrologic alteration.
43
 
h. The extraction of variables was executed with the Sample tool for obtaining the
final data matrix with the 11 independent variables and presence/absence of each
habitat type for total sample points.
44
 
Channel (CHAN)
Roads (ROAD)
River channel (RC)
Lagoons (LAG)
Buildings (BUI)
Constraining layer
Delta
Create random points
model_points
n = 1000
absence_points
n points = a
absence_points
absence_points
absence_points
absence_points
Coastal lagoon pres map
Tidal flat pres map
Salicornia-type pres maps
Salt meadows pres maps
…Rice fields pres maps
Create random points
n = a
absence_points
absence_points
absence_points
absence_points
presence_points
Erase
Rice fields (RICE)
RC
LAG
BAY
IB
OC
FR
RM
ROAD
CHANN
RICE
RC
LAG
BAY
IB
OC
FR
RM
ROAD
CHANN
RICE
Z
Coastal lagoon
Tidal flat
Salicornia-type
Salt meadows
Rice fields
…
Data matrix
for each
habitat type
Sample
Sample
Euclidean distance
1x1m
Fig. 19. Cartographic
model to generate
sample points and
extract the metrics
for predictive
wetland model
45
 
3.7 Statistical analysis
For each wetland habitat type, the presence/absence points and its respective terrain metrics
(elevation, distances to river/sea influence and distances to hydrologic alteration sources) was
determined with GIS-approach as described in the previous sections. PCA analysis was
initially performed to study the overall relations between elevation and distance variables.
Regression scores were extracted to analyze each wetland habitat pattern. Moreover,
differences on variables between wetland habitats were assessed for achieving their ranges
into the hydrological influence and soil altitude gradients. All the environmental independent
variables were checked previously for normality and linearity. All the variables required
natural logarithmic transformation to meet these parametric assumptions. Differences on
vertical soil elevation between habitats assessed by CORINE land cover and mean transects
was tested by Mann-Whitney test. Correlation analysis between elevation metrics (min, max,
range and mean) and distances was performed to check the independence of the variables. All
the statistical analyses were performed using SPSS v18.0 for Windows package.
Logistic multiple regression predictive model
As habitat map points are categorical variables (presence or absence), logistic regression is
proposed for predicting occurrence of wetland habitat types from topographical deltaic
variables (independent variables)(Hosmer and Lemeshow 2000). LMR is adequate because
the dependent variable is dichotomous (presence/absence) and the model admits non-
Gaussian independent variables (Franklin 1995). As explained, several authors proposed to
use equal number of presence and absence points for developing logistic method to avoid bias
(Felicísimo 2003; Narumalani et al. 1997). The foundation of the method is based on the
probability of occurrence of any number of classes of a dependent variable (in this study,
wetland habitat) based on explanatory variables (i.e elevation and distances). According to
Custer and Eveleigh (1986), “regression modeling involves the derivation of a mathematical
relationship between a set of independent predictor variables and a specific dependent
condition”. A multiple logistic regression analysis within the sample points generated in
previous steps resulted in an individual regression equation for each wetland habitat type.
46
 
Logistic regression is sensitive to extremely high correlations between variables that are
supposed to be independent (Hosmer and Lemeshow 2000). To avoid this, high correlation
between variables was eliminated by retaining only the variable with the highest explanatory
power for pairs of variables with r Pearson coefficient (r > 0,6)(Filipe et al. 2002). In the
multivariate analysis a forward stepwise was applied to each selected variable with a
probability of entry of 0,05 and removal of 0,10. The addition and exclusion of variables was
based on Wald’s test and the assessment of correlation was based on differences in the
coefficients estimated when a variables is added to the model and from partial correlation of
the estimated coefficients for p<0,001 (Zar 1999).
To assess the fit of each model, the Chi-square test and a classification table was used. This
procedure has been used by other authors (Narumalani et al. 1997; Ríos et al. 2005). The fit
test examined the deviance of the model with the constant versus the final model and rejection
was at p<0,05 significance based on a chi-square distribution. Another technique has been
used to evaluate each predictive model. The Receiver Operating Characteristic (ROC)
provides a threshold independent measure of accuracy and results in a plot of the relative
proportions of correctly classified sites over the whole range of threshold levels (Narumalani
et al. 1997; Tarkesh and Jetschke 2012). The ROC plot is obtained by plotting the sensitivity
of the model against the false positive fraction over all thresholds, being the area under the
curve (AUC) the probability that the model will distinguish correctly between observations.
An area of 1 is perfectly accurate, whereas on of 0,5 is performing a random model (no
acceptable). This fitting method has been applied in several studies of predictive vegetation
modelling (Álvarez-Arbesú and Felicísimo 2002; Felicísimo et al. 2002; Syphard and
Franklin 2010). All the statistical analyses were performed using SPSS v18.0 for Windows
package.
Model validation
The predictive model has been validated with independent data obtained by field surveys, in
which information about the presence/absence of different habitats was obtained by mean of
transects. So, the validation was applied to Salicornia marshes, salt meadows and reed beds.
47
 
3.8 Model implementation in the GIS
Logistic regression was used to generate probability models of habitat distribution with the
introduction of a spatial component through GIS. Several authors have already applied it to
coastal ecosystems (Álvarez-Arbesú and Felicísimo 2002; Narumalani et al. 1997; van
Horssen et al. 1999; vandeRijt et al. 1996) and other fields, such as forested areas (Felicísimo
et al. 2002; Turner et al. 2004) or soil mapping (Giasson et al. 2006). The LMR technique
yields coefficients for each variable based on data derived from samples taken across a study
site. These coefficients serve as weights in an algorithm which can be used in the GIS
database to produce a map depicting the probability of wetland habitats. Quantitatively, the
relationship between the "occurrence habitat" and its dependency on topographic variables
can be expressed as:
( ))()(...)1()1()0(
1
1
)( nxnbxbb
e
iP ⋅++⋅+−
+
=
where P(i) represent the probability value, x(1) … x(n) the values of the terrain variables and
b(1) … b(n) are coefficients derived from logistic regression. Each regression equation results
in a response value on an interval scale between 0 and 1. These response values can be
interpreted as a relative frequency of occurrence or an estimate of probability of occurrences
of wetland habitats. Then, low responses values will indicate low relative occurrence while
high response values indicate high relative occurrence. The probability of occurrence was
calculated from the logistic regression models in the raster calculator of ArcGIS.
The coefficients of terrain variables which resulted statistical significant in each logistic
model was applied to the GIS of Ebro Delta to produce a probability map of occurrence (cell
size = 1m.) for two wetland habitats: Salicornia-type marshes and salt meadows.
48
 
4. Results and discussion
4.1 Current distribution of wetland habitats in the Ebro Delta
Each wetland habitat was mapped and its extension was extracted in order to provide a
baseline data for wetland cover in the study area. The analysis and elaboration of these maps
through the existing CORINE land cover maps provided the more representative wetland
covers in the Ebro Delta. A total of nine habitats were analyzed by an elevation dependent-
approach, distance to river/influence and distance to sources of hydrologic alterations sources.
Table 5 presents the general descriptors about habitat occupation in the delta:
Table 5. Wetland habitat area in the Ebro Delta and its corresponding relative occupation. Urban areas and crops
other than rice are excluded. *Priority habitat. Surface area of Delta = 330 km2
(Ibàñez et al. 2010).
 
Habitat EU code Patch number Area (km2) % Habitats % Delta
Coastal lagoons *1150 34 16,85 5,86 5,11
Sandy habitats
2110,
2120,
2210
27 12,31 4,28 3,73
Tidal flats 1140 8 13,08 4,55 3,96
Salicornia-type marshes 1410 38 10,65 3,70 3,23
Salt meadows 1420 2 0,52 0,18 0,16
Cladium marshes *7210 7 3,35 1,17 1,02
Reed beds 53.1 29 8,67 3,01 2,63
Rice fields 82.d 10 222,05 77,17 67,29
Riparian vegetation 92A0 1 0,23 0,08 0,07
Total 287,75
Total surface occupied by wetland habitats was 287,75 km2
. Overall, rice fields are the
dominant habitat in the deltaic plain in terms of % coverage of the study area. Sandy habitats
(dunes and beaches) and tidal flats occupied near 10 % of the delta habitat surface, followed
by salt marshes with Salicornia and Juncus (4%). Helophytic habitats, represented in the
Delta principally by fresh and brackish marshes Cladium-type and reed beds, occupy 1,17 and
3,01% respectively.
49
 
Phragmites marshes (reed beds) have an important representation in the deltaic landscape
since occupy brackish (Garxal), fresh marshes (Vilacoto) and the altered margins of the salt
marshes due to agricultural runoff (Encanyissada, Aufacada). This habitat forms a great
diversity of plants associations from a physiognomic point of view due to high variability of
flooding levels and water salinity in the Delta (Curcó 2001).
Under natural conditions (i.e peat soils largely flooded by carbonate fresh waters) Cladium-
type marshes is limited to the natural wells “ullals” and some coastal lagoons that receives a
significant of fresh groundwater supply (Encanyissada, Vilacoto). The potential area of
Cladium marshes havs been modified by human activities (mainly agriculture) and its
hydrological pattern has been altered by the establishment of an extensive draining system to
lower the underground water level (Capítulo et al. 1994). This habitat is included in Annex 1
of the European Union Directive as a priority habitat type (7210 Calcareous fens with
Cladium mariscus and species of the Caricion davallianae).
The aquatic habitat of coastal lagoons is the other European priority habitat type included in
our study. Coastal lagoons of the Ebro Delta are the most representative aquatic habitats and
they occupy a higher surface area than the terrestrial habitats such as sand-dunes systems or
Phragmites beds. Their hydrologic conditions have changed over the past 100 years due to the
increment of the fresh water inputs and according Curcó and col. (1995) former coastal
lagoons occupied most their potential area and were more numerous. There is a high
variability in terms of surface of the lagoons: ranging from Garxal lagoon with 235 ha to
Encanyissada lagoon with 786 ha. The main source of variation in the coastal lagoons is their
hydrological regime which varies according to the fresh and salt water inputs. The freshest are
the west basin of the Encanyissada lagoon, les Olles and those having more connection with
the river (Garxal lagoon). The saltiest lagoons (la Tancada, Canal Vell or Buda), as other
Mediterranean coastal lagoons, have a hydrological pattern linked with seawater fluctuations
and some of them have periods of hypersalinity in summer (Badosa et al. 2006; Pérez-Ruzafa
et al. 2005).
50
 
Within the elevation gradient, the area occupied by wetland habitats in the 0,0 – 0,3 m range
is 73.54 km2
(26% of study area). From 0,3 to 0,5 m the surface occupied by wetlands is less
(25.31 km2
, 9%). The habitats surface between 0,5 to 1 m have been 7,5 km2
(2.6% of study
area). But between 1 and 1,5 meters of topographic elevation is where the largest area of
habitats is concentrated due to the presence of major part of rice fields in this elevation range
(168,22 km2
, 58%). The wetland area located beyond 1,5 meters only represents 0,31% of the
study area, being the sand-dune systems the more representative habitats in this elevation
range. The wetland habitats with mean elevations under mean sea level (-0,5 – 0,0 meters )
represent the 4,3 % of the total area (12,31 km2
). In this range we find mainly coastal lagoons,
tidal flats and reed beds. Table 6 summarizes the descriptive metrics of surface elevation for
each habitat type included in the study:
Table 6. Surface elevation metrics (min, max, rang and mean) for each wetland habitat. Values are
shown in median and SE (in brackets); n= number of polygons.
Habitat type n Min Max Range Mean
Coastal lagoons 34 -0,325
(0,055)
1,718
(0,138)
2,044
(0,146)
0,181
(0,038)
Sandy habitats 27 -0,145
(0,034)
2,731
(0,239)
2,875
(0,248)
0,698
(0,066)
Tidal flats 8 -0,314
(0,072)
1,718
(0,227)
2,031
(0,259)
0,227
(0,089)
Salicornia-type
marshes
38 -0,205
(0,052)
2,204
(0,156)
2,409
(0,167)
0,509
(0,036)
Salt meadows 2 0,410
(0,640)
2,275
(0,255)
1,865
(0,385)
1,125
(0,359)
Cladium marshes 7 -0,302
(0,171)
1,692
(0,292)
1,994
(0,356)
0,387
(0,134)
Reed beds 29 -0,690
(0,087)
1,900
(0,140)
2,590
(0,129)
0,301
(0,042)
Rice fields 10 -0,623
(0,194)
3,625
(0,582)
4,249
(0,628)
0,820
(0,255)
Riparian vegetation 1 0,490 4,230 3,740 2,808
Comparing the distribution of the eleven types of wetland habitats with the surface elevation
of the Ebro Delta, we conclude that the existence of zonation is low but differences were
found between habitats (Figure 20). One-way ANOVA tests (Table) indicate that there are
51
 
significant differences between the surface elevation of the wetland habitats observed.
Elevation metrics (min, max and mean) were previously log-transformed for achieving
parametric assumptions.
A B
C
Reedbeds
Ricefields
Lagoons
Tidalflats
Cladium
Salicornia
Sand-dune
Saltmeadows
Riparian
Reedbeds
Ricefields
Lagoons
Tidalflats
Cladium
Salicornia
Sand-dune
Riparian
Saltmeadows
Cladium
Reedbeds
Tidalflats
Lagoons
Salicornia
Salt
meadows
Ricefields
Riparian
Sand-dune
-1,000
0,000
1,000
2,000
a
b
a,b a,b a,b
a,b
a,b a,b
a a
a
a,b
a,b
a,b
a,b
b
a
a a,b
a,b a,b
b,c
b,c
c
1,000
2,000
3,000
4,000
5,000
0,000
0,500
1,000
1,500
2,000
2,500
3,000
Fig. 20. Surface elevation metrics (in meters) for each habitat type of the Ebro Delta. A: minimum, B:
maximum, C: mean. The error bars represents the standard error considering all the polygons of the
CORINE habitat map. Different letters point to significant differences (post hoc Tukey test: p < 0.05)
 
52
 
Table 7. One-way ANOVA results for the surface elevation metrics extracted from CORINE land
cover maps. Riparian vegetation was not included in the ANOVA analysis due to an insufficient
number of samples.
Elevation metrics
Sum of
Squares df Mean Square F Sig.
Minimum Between Groups 2,219 8 0,277 5,652 <0,000
Within Groups 7,163 146 0,049
Total 9,382 154
Mean Between Groups 0,465 8 0,058 13,408 <0,000
Within Groups 0,637 147 0,004
Total 1,101 155
Maximum Between Groups 1,113 8 0,139 5,566 <0,000
Within Groups 3,676 147 0,025
Total 4,789 155
Range Between Groups 0,873 8 0,109 4,387 <0,000
Within Groups 3,655 147 ,025
Total 4,528 155
Lower elevations, excluding coastal lagoons, are occupied more frequency by tidal flats and
reed bed habitats. Phragmites marshes are present in the lowest mean elevation in contact
directly with coastal lagoons (Fig. 20A), as mapped in the CORINE land cover. Even though,
the minimum elevation of emergent reed beds (-0,69 m.) has to taking into account since the
growing of these communities is limited in waters of 0,3 – 0,4 m. depth (Coops et al. 1996;
Squires and Valk 1992). Although scarce in the Delta (Ibàñez et al. 2002) this habitat type
will be present in permanent or nearly flooded soils of fresh-brackish marshes.
In salty coastal environments like the Ebro Delta, salt marshes are dominated by Salicornia-
type vegetation and depending on its relative soil elevation, different genera can dominate
(Ibàñez et al. 2010). In this study, Salicornia-type habitat had a mean elevation of 0,51 ± 0,04
m. Other authors (Pont et al. 2002) have found a similar topographic distribution of these
habitats in the Rhône Delta, which range between 0,25 and 0,60 m.
A more clear variation of soil elevation at upper regions was detected between Salicornia-
type habitat and salt meadows dominated by Juncus. Even though this halophytic habitat has
been drastically reduced in the Ebro Delta, its still occupies a broad range of elevation.
Regarding salinity, salt meadows with Juncus maritimus and Juncus acutus occur in soils less
53
 
influenced by the underground sea water level in contrast to the exclusive halophytic
communities (fruticose salt-marshes) (Espinar 2009). Thus, significant differences in mean
soil elevation of Salicornia-type and Juncus-type environments were found (unpaired t test, p
< 0,05). Silvestri and col. (2005) found a mean elevation difference between Juncus genus
and Arthrocnemum genus of 15 cm. Our results show that given habitats are found at higher
differentiated topographic elevations.
The riparian habitat was observed at the highest mean and maximum surface elevation of the
habitats (Fig. 20.D). The only patch mapped from CORINE land cover (Sapinya Island, 24
ha) shows a mean surface elevation of 2,81 m and its elevation can be considered
representative of this habitat. This habitat is present in the fluvial levees that are the highest
areas of the Delta plain, where it should develop according to the lowest salinity levels and
eventual flooding events. Presently, the potential area of riparian vegetation of the Delta,
especially Populus and Salix genus, has been transformed into rice fields and other crops.
The humanized habitat of rice fields is present in the maximum range soil elevation extracted
by the DEM of Deltaic plain. (Fig. 20.C). While other habitat types have a narrow elevation
range (i.e tidal flats or Cladium marshes) rice fields exhibit a wide distribution, indicating a
relative indifference to soil elevation. The high surface area occupied by rice throughout the
Delta plain, from near-river lévées to the margins of the coastal lagoons has led to major part
of the topographic gradient being occupied by this habitat. Moreover, the soil elevation of the
Delta has been altered in many areas by agricultural purposes, lowering the upper zones and
filling depressions. Then, the wide range in the elevation of this habitat could be attributed to
this human factor.
Methodological constrains on DEM application
In several habitats, no consistent results have been detected in the application of high precise
Digital Elevation Model of the Ebro Delta. Coastal lagoons showed mean elevation above sea
level (0,18 m.), and its maximum elevation was placed around 1,7 m. When extracting
elevation, the presence of micro-topography like “tores” (accumulation of soil in inundation
areas that it elevates above water surface) probably result in a bias of elevation values. The
level of detail in mean elevation of tidal flats (0,23 m.) it may associated to the same issue.
54
 
Depending on tidal range, this habitat should be located almost at sea level water or below sea
level (e.g. -0,3 m.)(Sakamaki et al. 2006). In contrast, the spatial resolution of the DEMs in
several studies where elevations are sampled at 30 m intervals, are more appropriate since its
extension of the study area varies over thousands of square kilometres. (Brown 1994).
Relationship between surface and soil elevation of wetland habitats
The figure (Figure 21) shows the area occupied by natural habitat types (except rice fields)
along the elevation gradient of deltaic plain. Habitats located in lowest elevations seem to be
associated with marine-influenced environments, which coastal lagoons and tidal flats have
maximum surface between 0,0 and 10 cm above mean sea level. Freshwater marsh habitats
such as Phragmites-type occurs in high frequency on 0-1 and 0-2 m soil elevation directly on
contact with water bodies that allow it flooded soils. Cladium marshes have showed a flat
surface distribution as evidenced by its minimum topographic position more elevated than
reed beds. Geographic position of Cladium habitats, near the inner border of the Delta
associated with continental groundwater discharge areas, could explain its surface elevation
pattern. Sandy habitats (dunes and beaches) are the habitat type with maximum surface at ca
0,5 m. Among this habitat, we can find several dune environments with different stability
stages. This succession pattern (i.e embryonic, shifting and fixed) largely determines their
topographic position along the elevation gradient of the Delta. Thus, according with the Delta
prevailing wind (NW), we can find different patterns of sand-dune surface occupation due to
the different orientation of the coast (Curcó 2006). At 0,6 – 0,7 m above the mean soil
elevation, a few patches of Juncus with relative surface overlap with Salicornia-type marshes,
especially in La Tancada. Finally, riparian vegetation has shown its maximum surface in the
highest zone, although its representation in fluvial lévées has decreased considerably.
55
 
Area(km2)
 
Fig. 21. Distribution of natural habitat types as function of soil elevation and surface. Rice fields were
not included in the plot.
56
 
Relationship between terrain variables
For the entire wetland habitats the lowest elevation, highest elevation, range elevation and
mean elevation show strong positive correlation (Pearson, p<0,01). The mean elevation has
been correlated positively with the distance to channels and negatively with distance to inner
border, lagoons, river and former river arms.
The three variables related with hydrologic alterations elements (roads, channels and rice
fields) were strongly (negatively) correlated to each other, which was expected due to their
overlapped position in the deltaic plain. Note that distance to former river arms of the Ebro
River (Riet de Zaida, Fondo and Muntells) was correlated positively with distance to rice
fields, channels and roads of the Delta. This can be explained because of these ancient courses
are largely occupied by rice fields and consequently by channels, either irrigation or drainage.
A significative (negative) correlation was found between distance to bay and river channel
that could represent a longitudinal gradient from fluvial lévées to bay (marine influence).
Distances associated with the riverine influence are also strongly correlated, mainly between
river channel and river mouth.
PCA ordination
In order to investigate the relations between all the environment variables on wetland habitats
distribution a principal component analysis (PCA) with varimax rotation was carried out
(Figure 22). Most of the analyzed variables were interdependent and have significative
correlation among them (Table correlations). The usefulness o the PCA was checked through
Kaiser-Meyer-Olkin’s (KMO) measure of adequacy sample (0,686) and Bartlett’s test of
sphericity (p<0,001). The two first axes explaining the 26% and 22% of the total variation
respectively. The minimum elevation of habitat polygons and distances to anthropogenic
elements was correlated positively with PCA axis 1. The first axis separated the patches of
wetland habitats closer to road, channels rice fields and former river arms with higher
minimum elevation than patches placed far of these elements (lower minimum elevations).
Then, the first axis summarizes the variation associated with rising elevation of wetland
habitat placed near anthropogenic elements and ancient river lévées. PCA axis 2 explains the
variation associated with maximum elevation gradient of habitat patches from interior to
exterior deltaic plain. The highest elevation of habitats patches has been correlated
57
 
(positively) with areas placed near the inner border of the Delta and opposites to exterior
limits of Delta (outer coast and river mouth).
The regression scores of each habitat polygon were extracted to visualize their position on the
two PCA axes. Analysis of salt meadows and riparian vegetation were not assessed due to its
lower polygons in the CORINE habitat map.
Some coastal lagoons were relatively separated on PCA axis 2: Buda lagoons were negatively
correlated with it (i.e. near from river mouth and outer coast and lower maximum elevations)
and patches of Encanyissada were correlated positively (i.e. far from river mouth and higher
maximum elevations).
Sand dunes showed patches separated along PC2. Sandy habitats correlated positively with
this axis have higher maximum elevations and are influenced by river mouth and outer coast
(e.g. Fangar dunes). While dunes/beaches oppositely placed along PCA axis 2 have shown
lower maximum elevations and were more influenced by bays and inner border (e.g.
dunes/beaches of Trabucador barrier). Tidal flats polygons have been differentiated in the
patches of Buda area (near river mouth and correlated positively with PC1) and patches of
Punta de la Banya.
The patches of Salicornia habitats were relatively separated along PCA axis 2. Then,
Salicornia-type marshes of Buda were different of polygons placed in la Banya given that the
last showed negative correlation with elevation and distance to river mouth. It means that
while there are some patches of Salicornia with lower maximum elevations located near the
bay, oppositely other groups with higher maximum elevations were placed near river mouth.
The reed beds were grouped according to PCA axis 2 mainly. Then, polygons with lower
maximum elevations and near to river mouth and outer coast were correlated negatively with
PC2 (i.e. reed beds of Buda lagoon as representative place). While patches of Phragmites
located in la Tancada seems to be associated with higher maximum elevations and greater
distances to these limits than either Buda or Garxal reed beds (positively correlated with
PC2). PCA axis 1 explains another source of variation related with reed beds distribution.
This axis separates groups of reed beds placed far from anthropogenic and former river arms
which can cause uprising elevation effect (higher minimum elevations).
58
 
The ordination technique has been useful to assess the relative distribution of wetland habitats
according to maximum variation sources of topographic deltaic variables. However, the
forcing factors of wetland distribution include other variables as soil salinity or moisture
content (Moffett et al. 2010). Regarding elevation, we can expect that habitats with higher
vertical elevation will have well-drained soils and lower flooding periods. Linking with the
soil salinity, the duration of evaporation periods (occurring when the marsh is not flooded)
increases with elevation and thus salts become increasingly concentrated (Adam 1993). Then,
stress conditions associated with salinity will be more evidence in lower regions within
intertidal range of the Ebro Delta (0-0,5 m)(Jiménez 1996). Soil salinity decreases beyond sea
water influence, therefore, these observations indirectly concerning the presence of wetland
habitats to topographic position of Delta.
59
 
 
Fig. 22. PCA-Ordination diagram of the environmental variables included in the study. For
abbreviations see the methods section.
60
 
Table 8. Mean soil elevation, distances to river/sea influence and distances to hydrological alteration of the main habitats in the Ebro Delta. Standard error of
mean in italics.
Habitat
Mean
elevation
(m)
Dist. to
outer
coast
(km)
Dist. to
inner
border
(km)
Dist. to
river
channel
(km)
Dist. to
lagoons
(km)
Dist. to
bay
(km)
Dist. to
river
mouth
(km)
Dist. to
former
river arms
(km)
Dist. to
rice
fields
(km)
Dist. to
channels
(km)
Dist. to
road
(km)
Coastal lagoons 0,181 1,549 11,381 6,409 0,000 3,933 13,078 6,278 1,517 0,141 0,085
0,038 0,344 0,801 0,953 0,000 0,607 1,511 0,743 0,456 0,048 0,031
Sandy habitats 0,698 0,656 8,325 9,265 1,046 1,608 16,406 7,401 2,825 0,977 0,758
0,066 0,160 0,842 1,073 0,210 0,441 1,905 1,136 0,503 0,246 0,244
Tidal flats 0,227 0,718 7,328 12,677 0,663 1,658 23,103 12,368 4,178 1,523 1,493
0,089 0,183 1,572 1,578 0,351 1,248 2,922 0,953 0,629 0,722 0,724
Salicornia-type 0,509 0,817 9,955 9,801 0,909 2,431 17,780 8,561 3,105 0,677 0,585
0,036 0,181 0,674 0,979 0,173 0,432 1,540 0,911 0,492 0,165 0,162
Salt meadows 1,125 4,559 7,301 4,237 0,502 2,916 18,096 4,489 0,000 0,000 0,000
0,359 4,342 6,829 0,231 0,135 0,884 6,676 3,851 0,000 0,000 0,000
Cladium-type 0,387 7,632 1,891 6,257 0,060 2,092 22,352 7,359 0,063 0,006 0,085
0,134 0,467 0,382 0,629 0,044 0,542 0,570 0,514 0,041 0,004 0,031
Reed bed 0,301 1,582 11,944 3,207 0,152 4,418 7,071 3,140 0,086 0,039 0,058
61
 
 
Table 8 continued
0,042 0,278 0,855 0,426 0,060 0,563 0,905 0,453 0,049 0,020 0,037
Rice fields 0,820 2,695 6,849 1,854 0,601 3,740 9,660 2,734 0,000 0,000 0,000
0,255 1,287 2,105 0,926 0,598 0,949 2,450 0,962 0,000 0,000 0,000
Riparian vegetation 2,808 10,537 3,674 0,000 6,215 8,458 18,067 1,686 0,201 0,166 0,055
0,000 0,000 0,000 0,000 0,000 0,000 0,000 0,000 0,000 0,000 0,000
ANOVA test p<0,0001 p<0,0001 p<0,0001 p<0,0001 p<0,0001 p<0,0001 p = 0,215 p<0,0001 p<0,0001 p<0,0001 p<0,0001
62
 
Table 9. Pearson’s correlation coefficients among independent variables for the current distribution of the habitats type in the Ebro Delta. Significance levels
** p<0,01; * p<0,05.
mean
ELEV
min
ELEV
max
ELEV
range
ELEV OC IB RC LAG BAY FR RM RICE CHANNEL ROAD
mean ELE 1
min ELE ,396**
1
max ELE ,618**
,110 1
range ELE ,374**
-,283**
,899**
1
OC -,067 -,054 -,287**
-,259**
1
IB -,323**
-,047 -,332**
-,288**
-,143 1
RC -,218**
,012 -,443**
-,423**
,233**
,014 1
LAG ,408**
,323**
,014 -,151 ,114 -,057 ,220**
1
BAY -,024 ,008 -,172*
-,189*
-,096 ,385**
-,275**
-,179*
1
FR -,198*
,215**
-,212**
-,314**
-,002 ,425**
,247**
-,021 ,105 1
RM ,017 ,027 -,280**
-,298**
,305**
-,196*
,698**
,281**
-,257**
,063 1
RICE ,050 ,423**
-,090 -,281**
,060 ,068 ,240**
,366**
-,188*
,474**
,135 1
CHANNEL ,184*
,416**
-,006 -,206*
-,001 ,036 ,211**
,298**
-,089 ,418**
,144 ,749**
1
ROAD ,030 ,359**
-,050 -,203*
,034 ,038 ,129 ,137 ,141 ,379**
-,003 ,605**
,703**
1
OC: Distance to outer coast
IB: Distance to inner border
RC: Distance to river channel
LAG: Distance to lagoons
BAY: Distance to bays
FR: Distance to former river arms
RM: Distance to river mouth
RICE: Distance to rice fields
CHANNEL: Distance to channels
ROAD: Distance to roads
63
 
Distances to riverine/marine and human infrastructures
Mean distances to river/sea influence (Figure 23) and human infraestructures (Figure
24) for each wetland habitat are plotted (Table 8). In general, the hydrological
boundaries associated with marine influence (the outer coast and bay mainly) have the
lowest distances to wetland habitats like tidal flats, sandy habitats (dunes and beaches)
and Salicornia-type marshes. Other habitats, such as Cladium-type marshes or riparian
vegetation showed higher distances to those variables (post hoc Tukey test p<0,05).
Phragmites marshes and salt meadows instead were located in intermediate distance
within this marine influence gradient. The effect of permanent flooded areas (i.e coastal
lagoons) has been demonstrated with the presence of emergent helophytic vegetation
(reed beds and Cladium marhes) placed closer than Juncus meadows. However, the few
samples cases of this habitat type (salt meadows) didn’t allow to assess the influence of
hydrological boundaries in a clear form. Except riparian vegetation, tidal flats, salt
marshes, sandy habitats and rice fields forms an homogenous group in relation with
distance to lagoons (Post hoc Tukey, p<0,05). That is, there are no statistical differences
in the distance to coastal lagoons of these wetlands habitats. These results confirm the
position of several habitats like patchwork of different classes without a clear pattern
around the lagoons.
Riverine influence
The geographic position of habitats according to its distance to inner border shows no
clear pattern, even though coastal habitats (i.e dunes and beaches, tidal flats), as
expected, were found at higher distances. The riverine influence expressed as distance
to river channel and river mouth mainly shows that riparian vegetation, reed beds, and
rice fields, in this order, are the closer habitats to these limits. The ANOVA tests
showed no significant differences in distance to river mouth of the wetland habitats (p =
0,215). A post hoc Tukey test indicated that the differences in mean distance to river
channel increases from reed beds and rice fields to all the other habitats. The same
results were found when the position of habitats respect to former river arms is
considered. In figure 23, we can observe increase in distance to riverine influences from
“fresher” habitats (closer) to marine habitats (far away). That is, tidal flats, Salicornia
64
 
marshes and sandy habitats were placed far away from river channel, river mouth and
former river arms.
Human infrastructures influence
Distances to hydrological alteration sources have shown significant differences of
wetland habitats studied (ANOVA test, p < 0,001 in all cases) (Figure 24). Reed beds,
salt meadows and Cladium marshes were the habitats closer to rice fields and channels
(mean distance < 100 m.) while Salicornia-type marshes, tidal flats and sandy habitats
were located far away (Post hoc Tukey test p<0,05). The close position of rice fields,
channels and roads to coastal lagoons (mean distance = 0,58 km), in contrast to other
habitats, such as Salicornia salt marshes (mean distance = 1,45 km) makes evidence
supporting the hypothesis of a hydrological alteration of closest alteration elements to
lagoons . The proximity of the disturbing elements can produce different effects
depending on their potential effects: hydrology balance between fresh (agricultural
runoff) and salt water, and barrier effect. Thus, effects of rice fields will base on the
variation of hydrologic balance between fresh water inputs during irrigation period and
salt water conditions during the rest of the year. Consequently, aquatic vegetation of
coastal lagoons is undergoing changes in their communities (Menéndez and Comin
2000; Menéndez et al. 2002). On the other hand, roads and tracks may act as barriers of
hydrological fluxes, and therefore, natural flows of water have been altered. The effects
of the proximity of channels on wetland distribution are difficult to discern using the
approach of the present study. Setting aside different hydrological effects (i.e drainage
or irrigation water), we can assume a barrier effect of these elements in the water natural
flow. So, the fact that the habitats are closer to hydrological alteration elements does not
mean a greater effect on their distribution.
65
 
0 2 4 6 8 10 12
Sandy habitats
Tidal flats
Salicornia-type
Coastal lagoons
Reed bed
Rice fields
Salt meadows
Cladium-type
Riparian vegetation
Distance to outer coast (km)
0 2 4 6 8 10 12 14 16
Rice fields
Riparian vegetation
Cladium-type
Reed bed
Salicornia-type
Tidal flats
Coastal lagoons
Sandy habitats
Salt meadows
Distance to inner border (km)
 
0 1 2 3 4 5 6 7 8 9
Sandy habitats
Tidal flats
Cladium-type
Salicornia-type
Salt meadows
Rice fields
Coastal lagoons
Reed bed
Riparian vegetation
Distance to bay (km)
 
0 1 2 3 4 5 6 7
Coastal lagoons
Cladium-type
Reed bed
Salt meadows
Rice fields
Tidal flats
Salicornia-type
Sandy habitats
Riparian vegetation
Distance to lagoons (km)
 
66
 
0 2 4 6 8 10 12 14 16
Riparian vegetation
Rice fields
Reed bed
Salt meadows
Cladium-type
Coastal lagoons
Sandy habitats
Salicornia-type
Tidal flats
Distance to river (km)
 
0 5 10 15 20 25 30
Reed bed
Rice fields
Coastal lagoons
Sandy habitats
Salicornia-type
Riparian vegetation
Salt meadows
Cladium-type
Tidal flats
Distance to river mouth (km)
 
0 2 4 6 8 10 12 14
Riparian vegetation
Rice fields
Reed bed
Salt meadows
Coastal lagoons
Cladium-type
Sandy habitats
Salicornia-type
Tidal flats
Distance to former river arms (km)
 
Fig. 23. Mean distances (in km) to river/sea influence for wetland habitats of the Ebro Delta. The errors bars represent the standard error of mean.
67
 
0 1 2 3 4 5 6
Salt meadows
Rice fields
Cladium-type
Reed bed
Riparian vegetation
Coastal lagoons
Sandy habitats
Salicornia-type
Tidal flats
Distance to rice fields (km)
 
0 1 1 2 2 3
Salt meadows
Rice fields
Cladium-type
Reed bed
Coastal lagoons
Riparian vegetation
Salicornia-type
Sandy habitats
Tidal flats
Distance to channels (km)
 
0 1 1 2 2 3
Salt meadows
Rice fields
Riparian vegetation
Reed bed
Coastal lagoons
Cladium-type
Salicornia-type
Sandy habitats
Tidal flats
Distance to roads (km)
 
Fig. 24. Mean distances (in km) to hydrological alterations for wetland habitats of the Ebro Delta. The errors bars represent the standard error of mean.
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Còpia de Benito_msthesis_FINAL2

  • 1. PREDICTIVE MODELLING OF WETLAND HABITATS IN THE EBRO DELTA WITH A GIS APPROACH         Xavier Benito Granell Màster en Planificació territorial: informació, eines i mètodes Facultat de Turisme i Geografia Universitat Rovira i Virgili IRTA – Unitat d’Ecosistemes Aquàtics
  • 2. 2   PREDICTIVE MODELLING OF WETLAND HABITATS IN THE EBRO DELTA WITH A GIS APPROACH Memòria del treball final del màster oficial de Planificació territorial: informació, eines i mètodes. Per: Xavier Benito Granell Dirigit per: Dr. Carles Ibàñez Martí Unitat d’Ecosistemes Aquàtics IRTA – Sant Carles de la Ràpita Dra. Rosa Trobajo Pujadas Unitat d’Ecosistemes Aquàtics IRTA – Sant Carles de la Ràpita Dra. Yolanda Pérez Albert Departament de Geografia Universitat Rovira i Virgili Vila-seca, Juliol de 2012
  • 3. 3   Agraïments Aquest treball ha estat possible gràcies una beca predoctoral de la Universitat Rovira i Virgili dins del conveni URV-IRTA. La base cartogràfica (Model digital d’elevació i ortofotomapes) són propietat de l’Institut Cartogràfic de Catalunya (www.icc.cat).
  • 4. 4   Table of contents Abstract ...................................................................................................................................... 5 1. Introduction............................................................................................................................ 6 1.1 General context: deltaic system, wetland habitats and human colonization .................... 6 1.2 The aquatic habitats of the Ebro Delta ............................................................................. 7 1.3 Predictive habitat modelling........................................................................................... 10 2. Hypotheses and objectives ................................................................................................... 15 3. Methods................................................................................................................................ 17 3.1 Study area ....................................................................................................................... 17 3.2 Wetland habitats, terrain variables and hydrologic alterations....................................... 19 3.3 Dependent variable: current distribution of wetland habitats......................................... 20 3.4 The independent variables: elevation and distances to hydrologic boundaries.............. 33 3.5 Vegetation transects........................................................................................................ 37 3.6 GIS development ............................................................................................................ 38 3.7 Statistical analysis........................................................................................................... 45 3.8 Model implementation in the GIS .................................................................................. 47 4. Results and discussion.......................................................................................................... 48 4.1 Current distribution of wetland habitats in the Ebro Delta............................................. 48 4.2 Logistic regression.......................................................................................................... 70 4.3 Probability of occurrence................................................................................................ 85 5. General conclusions ............................................................................................................. 91 6. References............................................................................................................................ 94  
  • 5. 5   Abstract Predictive habitat distribution models, derived by combining multivariate statistical methods with Geographical Information System (GIS) techniques, have been recognised for their utility in ecological modelling. The knowledge about potential distribution of natural habitats requires the link between current presence or absence of biological communities and a set of relevant environmental variables. This work examines the feasibility of using multiple logistic regression to model wetland habitat distribution in the Ebro Delta and to analyse how the riverine and marine influences affect its presence. Moreover, due to the high human occupation in the Delta, their influence in terms of distances to hydrologic alteration was assessed too. The predicted distribution was validated by comparison with a map of actual habitat type distribution (CORINE land cover) and by field transects. The variables that best explained the probability occurrence of habitats were soil elevation in habitats with higher mean elevation (e.g. Cladium-type marshes, dunes/beaches, rice fields and riparian vegetation) and distance to outer coast in habitats with lower elevations (coastal lagoons, tidal flats, Salicornia-type marshes and reed beds). The influence of the road and channels on habitats was reflected in higher soil elevations. The obtained prediction maps have provided the first results on habitat modelling in the Ebro Delta. The restricted distribution of some habitats due to human alteration may be the main reason of the mismatch between model predictions and field data in some habitats.
  • 6. 6   1. Introduction 1.1 General context: deltaic system, wetland habitats and human colonization The Ebro Delta (Catalonia, NE Iberian Peninsula) presents a formidable example of coastal wetland with a high variability of ecological factors (topography, edaphology, hydrology and climate) that play a key role in the configuration of its ecological gradients. The confluence of of contrasted dynamics (i.e riverine, marine and underground) explains most of the Deltaic variability across spatial and temporal scales. Due to human colonization (settlements, agriculture, hunting, turism...) wetland habitats with high natural value have been displaced in the periphery of the Delta where, significant remains of natural ecosystems subsist. In natural habitats whose sustainability and multi-functional values are threatened, like deltas, changes in land uses can have more environmental and ecological consequences than in other ecosystem. Deltaic ecosystems of the Ebro River have particular ecological and economical value because of their geographic position (interface between terrestrial and coastal zones) and diversity of habitats (wetlands, coastal lagoons, bays...) (Ibàñez et al. 2010). Human activities have led to a rapid deterioration of natural aquatic habitats since the beginning of XX century, mainly, owing to rice cultivation. Until then, however, the dominant landscape was determined by climatic, geomorphologic and biological factors, except areas transformed to saltworks. Due to its natural evolution, the Ebro Delta underwent geomorphic changes such as changes in river mouth and consequent erosion of abandoned lobes, filling of wetlands, accretion and subsidence of the deltaic plain or regression and coastal progression (Curcó et al. 1995). These natural processes led rapid modifications of the Delta’s configuration and in its ecological condition. After several millennia of growth, Ebro Delta was under a river- dominated dynamics but this trend changed a few decades ago in such a way that the present Delta is now a wave-dominated coast (Jiménez and Sánchez-Arcilla 1993). This change is mainly due to the construction of several dams along the river which has caused a nearly total reduction (97%) of the solid river discharge (Rovira and Ibàñez 2007). Overall, the drastic reduction of the sediment load slowed down the delta protrusion and intensified the delta coastline washout (Mikhailova 2003).
  • 7. 7   The Ebro Delta contains some of the most important wetland areas in the western Mediterranean. The majority of the Deltaic plain is devoted to rice agriculture and natural areas cover only about 25% of the total surface. Among these areas, a set of natural habitats are present: salt marshes, fresh-brackish marshes, coastal lagoons, sand dunes, natural springs and bays. Moreover, most of these habitats are included and protected by several European Directives (e.eg. Habitat Directive and Bird Directive) and regional laws (Ebro Delta Natural Park). This ecological diversity coexists with a human population near 50.000 inhabitants, which is located inside the Delta (15.000 inhabitants both Deltebre and Sant Jaume d’Enveja villages) and along the inner border (approximately 35.000 inhabitants with Amposta and Sant Carles de la Rapita villages). Urban zones, rice fields and other crops represents near 80% of total Delta surface. Intentisive human colonization in the Delta began at 1860’s with the first marsh transformations to rice fields after the construction of irrigation channels (southern hemidelta at 1860 and northern at 1912). From the beginning of 20th century until present, human transformation of the Ebro Delta largely occurred through draining of wetlands and the construction of an intensive irrigation system to bring fresh water from the river to rice fields. According to several authors (Curcó 2006; Mañosa et al. 2001) natural habitats declined its surface from 27.000 ha (80%) to 11.000 ha (30%) during the 1910-1960 period. The loss of natural habitats stopped during 1960s, but from 1970 to 1990 another 3000 ha of natural habitats were lost, leaving about 25% of total surface still occupied by lagoons and marshes. Such habitat loss has produced a change in the vegetal and animal communities present in the Delta. Present management in the Ebro Delta aims at maintaining a high agricultural productivity and valuable bird populations in the natural areas which are included in the Natural Park (Ibànez et al. 1997). 1.2 The aquatic habitats of the Ebro Delta The ecological value of the Delta reflects a high biodiversity, being the aquatic ecosystems the most important environments which support a representative sample of coastal wetlands. The Ebro Delta shows 18 natural habitats listed in the annex 1 of the European Directive on the conservation of natural habitats and of wild fauna and flora (Communities 1991). Bird and fish populations represent the major faunal groups in order of importance together with
  • 8. 8   singularity of halophilous and psammophilous plant communities. There are several technical reports (Curcó 2006) and scientific studies (Camp and Delgado 1987; Ibàñez et al. 1997; Menéndez et al. 2002; Valdemoro et al. 2007) that attempt to describe the aquatic environments of the Ebro Delta for achieving ecological information on their functioning. These studies note that there is a combination of habitats along a gradient of riverine and marine influence which confer wide environmental gradients. We consider the next main habitat units that coexist in the Deltaic plain: estuary, rice fields, coastal lagoons, natural wells, marshes, dunes and beaches, saltworks, bays and nearshore open sea domain. Estuaries are dynamic ecosystems that form a transition zone between river environments and ocean environments. Thus, estuaries are subjected to both marine influences (tides, influx of saline water...) and river influences (flows, topography of bed...). The pattern of dilution varies among systems. The Ebro Estuary is a salt-wedge estuary mainly dependent on the river discharge since the tidal amplitude range is very low (Ibàñez et al. 1997). Ebro Estuary extends from the mouth to 30 km approximately upstream, the position and presence of the salt wedge being determined by the tidal range and river discharge. The rice fields are the dominant landscape of the Delta and all aquatic ecosystems are influenced by water coming from rice fields. The hydroperiod associated with rice production is as follows: from April to December, a quantity of ca. 45 m3/s of river water is diverted to the irrigation canals for continuous irrigation. The resulting eutrophication associated with large amounts of fertilizer to enhance rice production, as well as pesticides, has led to a decrease in biological diversity and negative effects on aquatic vegetation of lagoons (Comín et al. 1991). Nevertheless, the rice fields form an aquatic matrix that link fluvial, lagoon and marine environments through network channels of irrigation and drainage. Coastal lagoons are littoral formations formed by the isolation of the marine domain through the development of a sand bar which separates the water bodies from the open sea (Kjerfve 1994). For their position and origin, their hydrological regime is determined by sea water inputs coupled with fresh water runoff from drainage of rice fields. For this reason, coastal lagoons of Ebro Delta show a hydrologic pattern that is clearly reversed (i.e hypersaline periods do not occur in summer as it would be expected in natural conditions). Aquatic macrophyte assemblages of the coastal lagoons (e.g. Buda lagoons) have been modified due
  • 9. 9   to the creation of salinity gradient that allows the development of different environments with the coexistence of several species of submerged macrophytes (Comín and Ferrer 1999). Marshes are wetland areas between terrestrial and marine domains which are linked with the coastal lagoons. Hydrological settings, mainly water and soil salinity, determine the presence of fresh (Vilacoto area), brakish (Garxal) and salt marshes (Buda Island) in the Ebro Delta. Vegetal communities of marshes are adapted to salinity content and soil moisture that generally depends on frequency and duration of flooding events (Bouma et al. 2005). Marshes play a key role exhibiting high primary productivity and assist important functions as nutrient removal and sediment retention (Ibàñez et al. 2002). Under global warming scenario that have produced eustatic sea level rise of 3.0 – 3,5 mmyr-1 over the past 15 years (IPCC 2001), scientific literature pointed out the ecological importance of marshes which retain sediments to offset sea level rise (Day et al. 2000). The natural freshwater wells are systems of natural ponds situated along the inner Delta border. In this area, underground water coming from often karstic inland areas (mainly Montsia) overflows through the surface forming small water bodies no more than 7 metres deep. A peat layer is present in this zone due to former palustral conditions, being catalogued as a priority habitat *7210 Calcareous fens in Council Directive 92/43/EEC. These freshwater wells are popularly called “ullals” and has been considerably altered by human activities, mainly through draining to lower the underground water level (Capítulo et al. 1994). As a result, Cladium marshes have been substituted by salt meadows dominated by Juncus genus in the area of “ullals” de Panxa. The shoreline of the Ebro Delta is occupied by sandy habitats that contain a very good representation of dunes and beaches. These habitats also exercise a key role in balancing the current coastline. The best dune systems of the Delta are placed in la Marquesa and el Fangar. According to soil mobility, there is a zonation from embryonic dunes (Agropyro-Honckenyion peploidis), shifting dunes with Ammophila arenaria and fixed dunes (Crucianellion maritimae). The development of tidal flats with microbial mats in the inner coast of la Banya peninsula represent another ecological value of these areas with particular hydrological conditions due to their differential orientation of its coast. Thus, lower topographic levels and NW dominant winds promote conditions to allow fluctuant moisture along drying and flooding events.
  • 10. 10   The bays are coastal marine waterbodies partially closed with a constant connection to the sea. In contrast to estuaries, the influence of freshwater is generally limited. In the Ebro Delta, Fangar and Alfacs bays were originated through the confinement of water bodies due the formation of spits parallel to the coast. The bays could be considered shallow coastal ecosystems (2 m. mean depth for Fangar bay and 3,13 m. mean depth for Alfacs bay)(Llebot et al. 2010), which involve marked spatial and temporal gradients. The saltworks are traditional salt production areas, located in zones where salt marshes should have their potential area. This artificial habitat is characterized by deep ponds with variable salinity that contributes to the increasing diversity of Delta. The Trinitat saltworks were included in the Natural Park since they provide favorable feeding habitats for the greater flamingo Phoenicopterus ruber, an emblematic species of the Ebro Delta. Nearshore open sea habitats are situated along the coastal area in front of the Ebro Delta. This environment borders on the whole deltaic plain, and the set of beaches, bars and spits are encompassed by it. A depth of 10 metres is considered the boundary between nearshore and offshore open sea. Nearshore waters can differ substantially from offshore waters due to the continental influence, and especially in an estuarine environment like delta. Thus, the latter are more eutrophic, with higher nutrient and chlorophyll concentrations and different phytoplankton composition (diatoms and dinoflagellates mainly). The high diversity of habitats and processes present in the Ebro Delta offers a unique opportunity to analyse the relationships between Ebro Delta habitats and their environment and to infer their potential distribution in relation to riverine and marine influences 1.3 Predictive habitat modelling The term “habitat” has been used in many ways in ecological studies. According to Spelleberg, (1994) habitat can defined as “the locality or area used by a population of organisms and the place where they live”. In ecology, the analysis of habitats-environment relationship has always been a central issue. The major factor involved in habitat distribution, especially in relation to plant communities, is climate in combination with geology or hydrology. Habitat factors that are playing a key role in species distribution should be
  • 11. 11   considered since species ranges and richness are often correlated with these factors (Vogiatzakis et al. 2006). The quantification of such relationships represents the core of predictive geographical modelling in ecology. These models are generally based on various hypotheses as to how environmental factors control the distribution of species and communities (Guisan and Zimmermann 2000). Hence, models of habitat distribution are not subjective models that predict how an area is suitable for development of a particular habitat in relation to environment conditions. The relationships between species, communtities or habitats (biotic entities) and environmental variables are frequently studied using gradient analysis that underlie hypotheses about species response functions (curves) to environmental gradients (Whittaker 1967). Austin and Smith (1989) defined three types of ecological gradients, namely indirect, direct and resource gradients. Indirect gradients have no direct physiological influence on species performance (slope, aspect, elevation, topographic position, geology). Direct gradients are environment parameters that have physiological importance, but are not consumed (e.g temperature, pH). Resource gradients address matter and energy consumed by plants or animals (nutrients, water, light, food for plants, water for animals…). Generally, literature pointed out that indirect variables usually replace a combination of different resources and direct gradients in a sample way (Guisan et al. 1998; Guisan et al. 1999). The real or actual vegetation is a patchwork of different classes or categories of communities. Classifying these communities in accordance with some key allows one to construct a vegetation map, which can be interpreted of actual vegetation that is normally more complex than habitat units map. While they are a simplification of reality, habitat maps are important data for correct environment management of a territory. The Ebro Delta has an important environmental dataset of which habitat maps are included. This work derived from the adaptation of the CORINE Biotopes project in Catalonia (Carreras and Diego 2007). But, this kind of information (i.e habitat maps or vegetation maps) has a limited temporal variability. Even in absence of human influence, vegetation dynamics is complex and intense, especially in deltas which are subject to sharp environment gradients. In the Ebro Delta, the ecological term of succession is applicable, defining the natural sequence in which a habitat replaces another over the passage of time. Then, the potential vegetation is defined as the stable community which would exist in an area as a consequence of progressive geobotanical succession if man ceased to affect and alter the terrain. Curcó et al. (1995) made an exercise
  • 12. 12   to delineate potential vegetal domains based on topographical and sedimentological features of the Deltaic plain. In this case, potential habitats will be more in balance with the salinity and moisture conditions of the environment. A model that considers sites with their disturbance features (e.g road or channels) might be expected to explain only a portion of the variance in habitat type distribution. Even so, this approach can be a chance for applying in the Delta. The first step that has to be considered in predictive modeling is the link between habitat units and mapped physical data. Several modelling methods have been used in scientific papers: heuristic, decision trees and statistical methods. The last approach, mainly regression, is the one most used to predict the value of the response variable if continuous, or the probability of a variable if categorical (Vogiatzakis 2003). Most predictive modelling efforts has used logistic regression to predict species (Rüger et al. 2005), vegetation assemblages (Davis and Goetz 1990) or animal habitat (Corsi et al. 1999). A logistic regression is well-suited where the dependent variable is dichotomous (presence/absence of habitats), and the technique allows one predictor (binary logistic regression) or more than one (multiple logistic regression). In addition the method lets a non-Gaussian distribution of the independent variables (Hosmer and Lemeshow 2000). Also, the result of the regressions ranges from 0 to 1 so that is appropriate for the generation of a likelihood model (Álvarez-Arbesú and Felicísimo 2002). The application of this method to wetlands and aquatic ecosystems is not an exception. (Narumalani et al. 1997) applied multiple logistic regression to predict aquatic macrophyte distribution. Another similar study was applied to aquatic vegetation by van de Rijt (1996) for predicting vegetation zonation in a former tidal area, while Shoutis et al. (2010) applied it for predicting riparian vegetation based on terrain variables and different river orders. The final step to consider in a predictive model is the model validation. Such evaluation consists in determining the suitability of a model for specific applications. According to Pearce and Ferrer (2000), wherever possible, evaluation is best undertaken with independent data collected form sites other than those used to develop the model. If independent data are not available, there are statistical techniques that fit the model in different degree, such as receiver operating characteristic (ROC) plot methodology (for more details see methods section). The next figure schematizes the generic steps of predictive modelling (Figure 1):
  • 13. 13   Fig. 1. Generic steps of predictive habitat modelling. The most used environmental predictors in predictive habitat modelling studies are those related with topography, geology and climate (Franklin 1995). Topography and its attributes such as elevation and slope, derived from Digital Elevation Model (DEM) are among the principal variables employed in these works due to their importance on vegetation patterns. A digital elevation model is any digital representation of the continuous variation of relief across space (Burrough et al. 1998). The use of accurate DEMs is especially important for deltas because, in the case of the Ebro Delta, about 40% of the plain surface lies under 0,5 meters above mean sea level (Ibàñez et al. 1996). In addition, hydrological variables such as frequency and duration of inundation are the main limiting factors of lagoon-wetland complex, and can be inferred through differences in soil elevation (Hickey and Bruce 2010). It is important to note that DEMs frequently contain systematic errors which can limit the effectiveness of predictive habitat distribution. On the other hand, too high accuracy will detect microtopography relief that may lead to unsatisfactory results. Ecological data sets have two distinct characteristics when compared to other kinds of data: they are multivariate and location specific. Recent studies to predictive modelling of habitats have been developed on a Geographic Information System (GIS), and ecological modellers have focus on incorporating spatial patterns in the models to apply them in large geographic areas (Vogiatzakis and Griffiths 2006; Zare Chahouki et al. 2010). Ecological modelling with GIS involves its complementary use for addressing ecological approaches, such potential distributions. In scientific literature, there are two ways of linking ecological models with GIS: 1) run the model outside the GIS and use the GIS for pre-processing data set (e.g. Statistical analysis Predictive modelling Model validation Environment Habitats
  • 14. 14   coordinate system transformation, location of sample points…) and generate cartographic outputs and 2) use GIS for extract metrics on environment mapped variables which will conforms the core of statistical method and post-processing of the data through cartographic display too (Felicísimo 2003; Felicísimo et al. 2002; Franklin et al. 2000). GIS-based spatial analysis tools facilitate the representation of ecological data across the space and its correlation with environmental data. In deltaic environment (among marshes, lagoons...), surface elevation (or water depth) and inundation frequency are the most important environment variables for vegetal zonation (Silvestri et al. 2005). Spatial analysis provides tools for researchers to assess how these factors influence habitat type distribution, extract metrics and explore the relationship between aquatic environments and topography by investigating species zonation. For example, Hickey et al. (2010) examined the relationship between distribution of salt marsh vegetation and the extent of tidal inundation using fine elevation data; (Xie et al. 2011) defined several landscape units of freshwater wetlands in Florida based on surface elevation; (Moran et al. 2008) linked spatial variation of flooding regime with the vegetation zones in a karst wetland. Habitats of the Ebro Delta are along environmental gradients which can be assessed to infer distribution patterns. Getting a predictive model, one can establish a relation between the habitat units and the environment data. The geographic scale of the Delta offers the opportunity to incorporate Geographic Information System techniques for extrapolating over a wide range those relations.    
  • 15. 15   2. Hypotheses and objectives Our initial hypotheses are: i) habitats of deltas are distributed spatially as a consequence of specific environmental requirements, mostly surface elevation and distance to sea/river. ii) These relationships (between delta habitats and surface elevation and the distance to sea/river) can be used to build a model that describes the potential distribution of each habitat according to the present configuration of deltaic plain. The main objective of this study was to determine the potential distribution of some existing wetland habitats in the Ebro Delta through a predictive habitat model based on terrain variables. To achieve this aim, the following specific objectives have been proposed: - To get elevation ranges of each habitat type within the altitude gradient of the delta by a digital elevation model (DEM). To validate them with field data. - To calculate distance ranges from the geographical position of each habitat type relative to the river and marine influence, which are determined from delta hydrologic boundaries. - To apply the predictive model in a Geographic Information System (GIS) to obtain maps of probability of presence for each habitat.
  • 16. 16   Research questions By achieving the objectives of the exercise, one would be able to answer the following questions: - What are the most important environment variables that determine the potential distribution of the main habitats in the Ebro Delta? - Are the human factors more important predictors than distances to hydrological boundaries in explaining the distribution patterns of wetland habitats? Or conversely, are the terrain variables? - The spatial distribution of aquatic habitats could be predicted accurately by developed predictive model? - Is the statistical technique of logistic regression appropriate to predict the link between physical variables and aquatic habitat types? - What are the ranges of elevation and distance to the hydrological boundaries of the delta of each habitat type? Research strategy This work is part of a broader study of palaeocological reconstruction of the Ebro Delta based on geochemical and biological analysis of its sediments. Two major steps can be distinguished within the research strategy of this Master Thesis: (1) modelling the link between the biological data and the accompanying terrain physical data and (2) implementation of the model in a Geographic Information System environment in order to attain a coverage habitat predictive map in terms of probability of occurrence for each aquatic habitat. These early steps will provide basic ecological information for evaluating the relationship between the types of aquatic habitats and the biological proxies (mainly fossil diatoms) preserved in its sediments.
  • 17. 17   3. Methods 3.1 Study area The study was carried out in the Ebro Delta, which is one of the largest deltas in the northwestern Mediterranean, with 330 km2 (Figure 2). Within this extension, rice fields occupy the majority of the delta plain (65% of the total surface) and natural areas cover only about 80 km2 (25%). These areas include a variety of aquatic habitats, providing an excellent example of coastal wetlands habitats: riparian vegetation, salt, brackish and fresh water marshes, coastal lagoons, natural springs, bays, sand dunes and mudflats. The confluence of contrasted dynamics, mainly riverine and marine, explains most of this high spatial variability. This diversity provides the presence of a large number of habitats of community interest listed in the annex 1 of the European Directive (Communities 2003) on the conservation of natural habitats and of wild fauna and flora. The best preserved natural areas are included in the Natural Park of the Ebro Delta that comprises 7.802 ha. Other zones that also include part of the rice fields are protected under other regulations of the Catalonian Government and the European Union (i.e Natura 2000).
  • 18. 18     Fig. 2. Location of the study area, the Ebro Delta.   There were various reasons for choosing this study area. Firstly, there is good information available, both biological (habitat maps) and physical data (digital elevation model). Secondly, there is a great spatial heterogeneity, favouring the existence of diverse environments, and allowing a wide range of ecological gradients to be assessed with the actual deltaic plain configuration. And thirdly, there is a wide set of aerial and topographic maps with different scales for carrying the GIS dataset analysis. The Ebro Delta shows a very low relief, with slopes of around 0,01 – 0,02 %. Riverine and sedimentary dynamics determine a elevation gradient decreasing from lévées in the inner border (4-4,5 m) to the river mouth (0-0,5 m) (Figure 3). Lévées of former river arms have more elevation than the adjacent deltaic plain and these structures can be recognised in the topographical maps. The low elevation areas are usually the ones having more marine influence. The agricultural activity has been the major factor modifying of the native
  • 19. 19   topography of the delta, either lowering high areas or filling lagoons. Moreover, currently the Ebro Delta is undergoing elevation loss due to sediment deficit created by sediment retention in the dam system along the river. This fact tends to lead a negative balance between vertical accretion and subsidence on in the deltaic plain. Furthermore, elevation loss is accelerated by sea level rise due to the effects of climate change. 300000 ,000000 300000 ,000000 320000 ,000000 320000 ,000000 4500000 ,000000 4500000 ,000000 Elevation (m) 0 - 0,5 0,5 - 1,0 1,0 - 1,5 1,5 - 2,0 2,0 - 2,5 2,5 - 3,0 3,0 - 3,5 3,5 - 4,0 > 4,00 5 102,5 Km. ´   Fig. 3. Digital Elevation Model of the Ebro Delta. Source: Cartographic Institute of Catalonia, 2011. 3.2 Wetland habitats, terrain variables and hydrologic alterations The natural habitat classification and mapping of the CORINE Biotopes (Communities 1991) developed for Catalonia was used to identify and select each wetland cover type. This data source includes two types of information, 1) the list of CORINE habitats of Catalonia (Vigo et al. 2006) and 2) the mapping of habitats in Catalonia 1:50,000 (Carreras and Diego 2007). The list has a hierarchical structure based on habitats classification of annex 1 of European Union Habitats Directive and describes each habitat unit from physiognomical, ecological and phytosociological characters. Overall, 9 habitat types have been selected to develop the model. The selection of wetland habitats responds to different criteria as a function of the
  • 20. 20   variability on hydrological requirements and salinity tolerances. Since the Ebro delta is a coastal system, the distribution patterns of broad types of wetland habitats, such as Salicornia-type marshes or salt meadows can be influenced by salinity. Without freshwater inputs, topography should be the main factor determining the habitat distribution. It is known that flooding regime is a primary factor structuring coastal wetlands with the frequency and duration of inundation determined by surface elevation (Hickey and Bruce 2010). In addition, the geographical position of each habitat respect fluvial and marine influence will affect its distribution in the deltaic landscape. In this study we have assessed how the target habitats are distributed through a combination of several distances that are related with the hydrologic boundaries of the delta plain (see section 3.5). However, the effects of the hydrologic alterations produced by two main anthropogenic sources should also be taken into account, these being 1) fresh water inputs due to irrigation from adjacent rice fields and the network irrigation channels; 2) roads that can interfere natural hydrologic fluxes. Thus, distances to these hydrologic alteration sources have been included as well in the model as possible predictors of geographic distribution of the wetlands habitats. 3.3 Dependent variable: current distribution of wetland habitats The presence or absence of the wetlands habitats has constituted the dependent variable of the model. For this purpose, the map of natural habitats on 1:50.000 scale has been used. This data set was acquired through digital format (shape file on ArcView environment) from the Environment Department of the Government of Catalonia. The sheets that cover the Ebro Delta include numerous habitats, from the dune domain to reed beds; at the same time each polygon comprises several classifications. In this study we chosed the main wetland habitats present in the Ebro Delta and the final delimitation of target habitats was subject to expert review. Most of the habitats (7 out 9, except reed beds and rice fields) are classified as community interest by the European Union Habitats Directive. The directive defines habitats of Interest as those that (i) are in danger of disappearance in their natural range; or (ii) have a small natural range following their regression or by reason of their intrinsically restricted area; or (iii) present outstanding examples of typical characteristics of one or more of the
  • 21. 21   seven following biogeographical regions: Alpine, Atlantic, Boreal, Continental, Macaronesian, Mediterranean and Pannonian. The Interpretation manual of European habitats (Romao 1996) was used to describe each wetland habitat from digital maps of the Ebro Delta. The list in table 1 shows the habitats included in the study and its corresponding classification according to CORINE Biotope classification, and it also lists the most representative sites of Delta where these habitats are present. So, this classification system has resulted in the list of habitats of Catalonia. The codification system of habitats is based on a hierarchical classification and has been identified by a code like nn.xxxx, where the first two digits indicate the main group it belongs to (Table 2). Thus, the code of each habitat provides information on the groups and subgroups to which they belong and with which other habitats have similarities. Table 1. Main CORINE groups of European habitats classification. Habitat CORINE group Coastal and halophytic communities 10 Non-marine waters 20 Shrubby vegetation and grassland 30 Forests 40 Bogs and marshes 50 Screes 60 Agricultural land and artificial landscapes 80 Burned areas 90
  • 22. 22   Table 2. Wetland habitats included in the study and its corresponding classification based on CORINE biotope project. * Priority habitat. Wetland habitat Code HCI CORINE code Delta sites 1. Coastal lagoons *1150 Coastal lagoons 21 Lagoons Encanyissada, Tancada, Aufacada, Platjola, Illa de Buda, Garxal, Canal Vell, les Olles 2. Sandy habitats 2110 Embryonic shifting dunes 2120 Shifting dunes along the shoreline with Ammophila arenaria (white dunes) 2210 Crucianellion maritimae fixed beach dunes 16.1 Sand beaches 16.2 Dunes Along shoreline of the Delta plain 3. Tidal flats 1140 Mudflats and sandflats not covered by seawater at low tide 14 Mud flats and sand flats La Banya, Fangar 4. Salicornia-type marshes 1420 Mediterranean and thermo-Atlantic halophilous scrubs (Sarcocornetea fruticosi) 15.6 Halophilous shrubby formations Buda island (Calaixos), Tancada 5. Salt meadows 1410 Mediterranean salt meadows (Juncetalia maritimi) 15.5 Mediterranean salt meadows Sant Antoni, Garxal, Tancada, Encanyissada 6. Cladium-type marshes *7210 Calcareous fens with Cladium mariscus 53.3 Cladium mariscus- dominated formations Vilacoto, Ullals of Baltassar 7. Reed beds - 53.1 Reed beds Garxal, Encanyissada, Tancada, Platjola, Aufacada, Canal Vell, les Olles 8. Rice fields - 82d Rice fields Over the deltaic plain except peripheral areas 9. Riparian vegetation 92A0 Salix alba and Populus alba galleries 44.1 Riparian willow formations Sapinya island
  • 23. 23   Legend Coastal lagoons (1150) Sandy habitats (dunes and beaches) (2110, 2120, 2210) Tidal flats (1140) Salicornia-type marshes (1420) Salt meadows (1410) Cladium fens (7210) Reed beds (53.1) Rice fields (82d) Riparian vegetation (92A0) Ebro river Human settlements0 5 102,5 km. ´ Fig. 4. Map of habitats of the Ebro Delta with its corresponding CORINE code
  • 24. 24   Description of wetland habitats 1. Coastal lagoons EU habitat code: *1150; CORINE code: 21 Coastal lagoons of the Ebro Delta are typical water bodies from a deltaic environment (albufera-type) formed due to the evolution of the Delta lobes. Because of their origin and separation from the open sea by a sand bar, they are lagoons strongly influenced by seawater inflows. In their original state they were salt water lagoons with a maximum salinity in summer, but due to the rice field drainage, their hydrologic regime has been severely. Coastal lagoons are zones with high biological and ornithological importance, where several species listed in the Bird European Directive are present. The aquatic vegetation of the coastal lagoons is composed of mixed macrophyte beds of Ruppia cirrhosa, Potamogeton pectinatus and Zostera sp. (Menéndez et al. 2002). There are a total of nine coastal lagoons in the delta, among them Buda island, placed near the river mouth, or the Aufacada lagoon.   Fig. 5. Present distribution of the coastal lagoons in the Ebro Delta according to the habitat mapping of Catalonia. Source: Department of Sustainability and Territory, Catalonia Government.
  • 25. 25   2. Sandy habitats (dunes and beaches) EU habitat code: 2110, 2120, 2210; CORINE code: 16.1, 16.2 These habitats are basically constituted by deltaic-front sand bodies. Ecologically sandy habitats can be characterized as environment that contents low water content, low levels of salts and organic matter and relative levels of mobility. Sandy habitats of the Ebro Delta bring together three types of habitats of community interest related to the substrate mobility: embryonic dunes (2110), shifting dunes with Ammophila arenaria (2120) and fixed dunes (2210). These habitat types are considerate as transitional and littoral sedimentary environments due to marine agents produce largely the mobilization of its soils. Then, they are associated with littoral transfer process. The extension of this habitat in both hemideltas is unequal. The main reason is the different orientation of the outer coast with respect to prevailing winds (NW). Thus, the most representative area of beaches and dunes systems is located in the northern hemidelta: Marquesa beach-Garxal and Punta del Fangar.   Fig. 6. Present distribution of the sandy habitats (dunes and beaches) in the Ebro Delta according to the habitat mapping of Catalonia. Source: Department of Sustainability and Territory, Catalonia Government.
  • 26. 26   3. Tidal flats  EU habitat code: 1140; CORINE code: 14 Flat coastal areas, devoid of terrestrial vascular plants and usually colonised by blue- green algae and diatoms. This habitat occupies coastal sands and muds and their associated coastal lagoons that experience recurrent episodes of flooding and drying. It is particularly well developed and forms the greatest extension in the Alfacs Peninsula, formed by la Banya spit and Trabucador barrier. This area is very sandy, and flooding periods are frequent due the strong northwestern winds, which results in a vertical stratification of physicochemical gradients between the aqueous interface and the solid substrate (Mir et al. 2000).   Fig. 7. Present distribution of the tidal flats in the Ebro Delta according to the habitat mapping of Catalonia. Source: Department of Sustainability and Territory, Catalonia Government.
  • 27. 27   4. Salicornia-type marshes EU Habitat code: 1420; CORINE code: 15.6 Low shrubby expanses of woody glassworts which in the Ebro Delta are dominated by succulent perennial species of the genus Sarcocornia and Arthrocnemum. Within the water salinity gradient of marshes, salt marshes are the wetlands with major influence of marine water. In them, the connexion to freshwater is limited, except for those zones that are receiving water inflows from of adjacent rice fields. Depending on rainfall, evaporation and tidal exchange, the salinity pattern may differ through the year. The differences of these factors can influence the ecological and physical traits of each marsh, such us vegetal communities (halophytic and hydrophytic), net primary productivity or accretion and subsidence rates. Buda Island is the most representative Arthrocnemum-type marsh in the Delta.   Fig. 8. Present distribution of the Salicornia-type marshes in the Ebro Delta according to the habitat mapping of Catalonia. Source: Department of Sustainability and Territory, Catalonia Government.
  • 28. 28   5. Salt meadows EU Habitat code: 1410; CORINE code: 15.5 This habitat is characterized by the presence of Juncus acutus and Juncus maritimus as the most representative plant. These taxa withstand high soil humidity and for this reason grow in drenched and/or periodically submersed soils. However, the habitat finds its ecological optimum in sites occurring at least a few centimetres higher than the average soil water level. In the Ebro Delta, it grows in scattered inland sites where soil elevation is higher than those of the halophilous scrub. Regarding salinity, this habitat forms a transitional stage between salt marshes Salicornia-type and habitats lacking halophytic vegetation. In the Ebro Delta, the salt meadows can form intermediate stands with halophilous scrubs. According to Curcó et al. (1995) this terrestrial habitat have been drastically reduced in relation to their potential surface area since that area has been impounded by the rice fields.   Fig. 9. Present distribution of the salt meadows in the Ebro Delta according to the habitat mapping of Catalonia. Source: Department of Sustainability and Territory, Catalonia Government.
  • 29. 29   6. Cladium-type marshes EU Habitat code: 7210; CORINE code: 16.2112 This habitat type is the only one the wetland habitats considered that constitute a priority habitat in the Ebro Delta. Within the fresh water ecosystems, the presence of Cladium-type marshes was originally associated with underground freshwater springs in karstic zones (Ullals) or in elevated zones with recurrent flooding events. Nowadays the most representative zone of this habitat in the Delta is in the Vilacoto area at the east of the Encanyissada lagoon. In this habitat the presence of dense helophytic communities dominated by Cladium mariscus, Phragmites australis and Scirpus maritimus is linked with a superficial peat layer and a significant input of underground water; this fact allow the submersion of the base of the plant during most of the year.   Fig. 10. Present distribution of the Cladium-type marshes in the Ebro Delta according to the habitat mapping of Catalonia. Source: Department of Sustainability and Territory, Catalonia Government.
  • 30. 30   7. Reed beds EU Habitat code: - ;CORINE code: 53.1 The habitat occurs near lagoons, channels or other wetland types which receive direct fresh water influence from the rice fields and the river. It occurs in still, fresh or brackish water. Within the Ebro Delta, natural colonies of Phragmites australis develop in the Garxal area, which is subjected to the direct influence of the riverine processes. Along the south edge of the lagoon there is an intermediate belt of brackish reedswamp dominated by Phragmites and Juncus species. Over the Delta plain, this habitat has spread along the margins of the coastal lagoons and bays due to hydrological changes caused by rice cultivation mainly.   Fig. 11. Present distribution of the reed beds in the Ebro Delta according to the habitat mapping of Catalonia. Source: Department of Sustainability and Territory, Catalonia Government.
  • 31. 31   8. Rice fields EU Habitat code:- ; CORINE code: 82d This habitat is the dominant landscape of the Delta as a result of a large agricultural occupation process that has led an actual coverage of near the 70% of the deltaic plain. Despite being a humanized environment and classified as artificial landscape for CORINE Biotope project, the rice fields constitute a aquatic matrix that link fluvial, lagoon and marine ecosystems. During the rice inundation period (May-December) this habitat acts as an authentic aquatic ecosystem which offers zones of feeding and resting to aquatic birds. Nevertheless, the inflow of huge amounts of fresh water into the fields has been an important factor in alterating the hydrology of the Delta, as well as causing loss of wetlands habitats and loss of elevation of the deltaic plain.   Fig. 12. Present distribution of the rice fields in the Ebro Delta according to the habitat mapping of Catalonia. Source: Department of Sustainability and Territory, Catalonia Government.
  • 32. 32   9. Riparian vegetation EU habitat code: 92A0; CORINE code: 44.1 This type of habitat is found growing in the Sapinya Island. According to CORINE land cover, this is the only patch of riparian vegetation present in the Ebro Delta. Under natural conditions, the lower Ebro River was bordered by riparian forest along its levees. Generally, this habitat types inhabited in the fluvial levees in mean elevation range of 2 and 4 m above sea water level. Flooding events in these areas only occurred when the river overflowed large flows, but due to construction of the dam system along Ebro river watershed the river flow has been drastically laminated. Coupled with human colonization of delta plain in terms of agricultural purposes, which was more significant in these higher zones, the riparian habitats have a relictual distribution.   Fig. 13. Present distribution of the riparian vegeation in the Ebro Delta according to the habitat mapping of Catalonia. Source: Department of Sustainability and Territory, Catalonia Government.
  • 33. 33   3.4 The independent variables: elevation and distances to hydrologic boundaries The Ebro Delta and the Mediterranean deltaic systems generally have a complex structure and its functioning depends on hydrologic, geologic and climatic factors. The diversity of habitats in the area of study is high, forming a set of environments that are river- and marine-dominated. The first factor has lost importance due to the dam construction along the Ebro River watershed concerning the reduction of near the 99% in the particulate sediments of the lower river. Today, the hydrological conditions in some salt marshes of the Delta are dominated by inputs of seawater through outlet channels, much more than riverine influence, being the agricultural runoff the factor that is altering the natural conditions (except in the river mouth area, Garxal). The topographical factor plays a key role in the Ebro Delta since about 40% of the plain surface lies under 0,5 meters above mean sea level (Ibàñez et al. 1996). In addition, hydrological factors are highly correlated with the variation of soil elevation that will determine frequency and duration of the inundation events. The distribution and composition of lagoon-marshes complexes depends strongly on this terrain variable. Thus, river lévées are the highest parts of the Delta, and under natural conditions, are vegetated by riparian forests such as Populus and Salix galleries. These habitats are flooded only during high discharge. Outside these areas are fresh, brackish or salt marshes, depending on factors such elevation, inputs of upland runoff, riverine influence, marine influence or soil drainage. Regarding vegetation marsh zonation, in some cases there is a clear vegetation transition related with soil salinity and water regime (Silvestri et al. 2005). The link between these terrain factors and vegetal communities is one of the research questions of this study. The independent variables included in this study for assessing the potential distribution of the wetland habitats have been surface elevation, distance to hydrologic alterations and distance to river/sea influence. The last approximation was assessed by the combination of several distances which are associated with the hydrologic boundaries of the Delta plain and will serve to extract influence of the flooding regime as an indirect way. The hydrologic alteration approach includes all of the elements on the deltaic
  • 34. 34   landscape that have resulted from human activity, mainly roads, irrigation channels and rice fields. Table 3. List of terrain predictors included in the study. Terrain variable Abbreviation Description Source Surface elevation ALT Terrain altitude of the delta plain Digital elevation model 1x1 m (Base cartogràfica de l’Institut Cartogràfic de Catalunya) Distances to river/sea influence - Outer coast OC External shoreline Topographical map 1, 25.000 (ICC) - Inner border IB Inner side of deltaic plain Topographical map and ortophotomaps (ICC) 1, 25.000 - River channel RC Ebro river course and its levees Topographical map 1, 25.000 (ICC) - Lagoon LAG Coastal lagoons Topographical map 1, 25.000 (ICC) - Bay BAY Coastal marine water bodies partially closed Topographical map 1, 25.000 (ICC) - Former river arms FR Ancient river courses: riet Fondo, riet de Zaida and riet Vell Digital elevation model (ICC) - River mouth RM Current mouth of the Ebro river Topographical map 1, 25.000 (ICC) Distances to hydrologic alteration - Roads ROAD Roads and paths constructed over the Delta plain Topographical map 1, 25.000 (ICC) - Channels CHAN Network of channels for irrigation and drainage waters from rice fields Topographical map 1, 25.000 (ICC) - Rice fields RF Agricultural crops of rice CORINE land cover (Department of Sustainability and Territory)
  • 35. 35   Legend Ebro river Coastal lagoons Bay Former river arms River mouth Outer coast Inner border0 5 102,5 km. ´ Fig. 14. Map of hydrologic boundaries from river/sea influences in the Ebro delta.
  • 36. 36   Legend Ebro river Coastal lagoons Channels Roads Rice fields5 0 52,5 km. ´ Fig. 15. Map of hydrologic alterations elements from human colonization in the Ebro delta.
  • 37. 37   3.5 Vegetation transects In order to validate the soil elevation of wetland habitats obtained via CORINE land cover, transects that cover soil elevation gradient has been developed by mean transects. Along transects, the presence of each habitat through the recognition of homogenous belts was recorded and sample points were georeferenced (European Datum 1950, UTM 31N). The next habitats were surveyed: Salicornia marshes, Juncus marshes and reed beds (fresh-brackish marsh) (Figure 16). Transects of salt marshes of Salicornia-type were developed in Sant Antoni Island, a marine-influenced area of Buda Island. In this site, succulent Salicornia and Juncus often co-occur. The Garxal brackish marsh bordered lagoon which receive river discharge directly. Transects were develop along south edge of the lagoon, where a belt of Phragmites marshes is present. The habitat of salt meadows dominated by Juncus genera was located in la Tancada area and la Platjola. Salt meadows of la Tancada are the area of Delta coincident with CORINE habitat map since in other marshes it were been detected (i.e. Garxal marsh) but it didn’t incorporated in the digital maps. Ebro river Coastal lagoons5 0 52,5 km. ´ Tancada: Juncus and Salicornia marshes Sant Antoni island: Salicornia and Juncus marshes Garxal marsh: Phragmites and Juncus marshes   Fig. 16. Location of study of marsh study sites in the Ebro Delta where elevation transects were developed.
  • 38. 38   3.6 GIS development Geographic Information Systems (GIS) are widely used for ecological studies because they provide techniques to relate several environmental/landscape variables and their specific location. This approach allows to assess the distribution of habitats according to environmental gradients and to suggest correlations between them. The first step has consisted to obtain the database that integrates the main variables included in the study: current distribution of wetland habitats and terrain deltaic predictors. Therefore the process was sequential: initially a database was established and then the model was applied for establishing the relationship between the wetland type distribution and terrain variables. A flow diagram of the general process is presented in Figure 17, and the specific cartographic methods will be explained in the following sections. Habitat maps Coastal lagoons, pres/abs Topographic maps Ortophotomaps Digital elevation model Distances to river/sea influence Distances to hydrologic alteration Terrain predictors Distribution of wetland habitats Tidal flats, pres/abs Wetland type, pres/abs . . . Logistic regression Predictive wetland habitat model Random sampling points, pres/abs Predictive habitat mapping Validation   Fig. 17. Flowchart of the process of generating the predictive model (grey boxes) and the current distribution of wetland habitat types of the Ebro Delta.
  • 39. 39   Database development Wetland habitat data were collected following a selection of habitat type layers using the CORINE land cover as vector format. A Geographical Information System (GIS) has been developed to generate the database of wetland habitat types (presence and absence) in the Ebro Delta. For this purpose, the commercial GIS package ARC/INFO, ArcView 9.3 was used. The cartographic data sources required to generate wetland covers and terrain variables are listed in the follow table (Table 4). All of the layers have been projected in UTM north zone 31, European Datum 50. Table 4. Data sources for obtaining digital database of wetland types and terrain variables.   Layer Format Scale/ Precision Output layers Source CORINE land cover Polygon, Shapefile (ArcView) 1, 50.000 - Pres/abs of each habitat type (n = 9) - Rice fields Department of Sustainability and territory (Government of Catalonia) Digital Elevation Model ASCII, raster (ArcView) 1x1 m - Raster elevation grid - Former river arms Cartographic Catalan Institute (ICC) Topographical map Polygon, Shapefile (ArcView) 1, 25.000 - Ebro river - Lagoons - Bays Cartographic Catalan Institute (ICC) Polygon, shapefile 1, 25.000 - Buildings Cartographic Catalan Institute (ICC) Polyline, shapefile 1, 25.000 - Outer coast - Irrigation channels - Roads Cartographic Catalan Institute (ICC) Ortophotomaps MrSID, raster 1, 5.000 - Deltaic plain - Inner border - River mouth Cartographic Catalan Institute (ICC)
  • 40. 40   The complete cartographic procedure for obtaining the database is shown in Figure 18, but the general steps can be summarized as follow: a. Digitalization of the Delta plain to obtain the geographical limit of the habitat and distance layers. b. Obtaining the habitat map cover for each target wetland (vectorial format): coastal lagoons, salt meadows, Salicornia-type marshes, Cladium marshes, Reed beds and rice fields. The habitat map cover of sandy habitats (dunes and beaches) and tidal flats were modified from the original sheets through expert criteria. c. Extraction from the topographic sheets (1,25.000 scale) the hydrologic layer in vectorial format (polygon and polyline). From the polygon data, select by attributes the cases of river courses (RC), lagoons (LAG) and marine waters (BAY). From the polyline data, select by attributes the cases of outer coast (OC) and channels (CHANN). d. Extraction from the topographic sheets (1,25.000 scale) the settlement layer in vectorial format (polygon and polyline). From the polygon data, select by attributes the cases of generic buildings (BUIL). From the polyline data, extract by attributes the cases of roads (ROAD). e. Digitalization of the inner border (IB) and river mouth (RM) in polyline format from the ortophotomaps on 1, 5.000 scale. f. Recognition and digitalization of the former river arms (FR) from the DEM of Delta. g. From each wetland habitat map polygons with habitat presence, elevation metrics (min, max, range and mean) were extracted with zonal statistics tool of the Analysis tools extension. h. From each wetland habitat map polygons with habitat presence, distances to closest feature of hydrologic and anthropogenic layers was calculated with near tool of the Analysis tools extension.
  • 41. 41   Ortophotomaps Habitat maps cover Raster data Vector data Inner border (IB) Digit. Digit. Topographic maps Hydrologic polygon layer River channel (RC) Channel (CHAN) Outer coast (OC) Lagoons (LAG) Bays (BAY) Select by attributes CAS = MAI003 CAS = MAI001 CAS = MAI011 Hydrologic polyline layer Select by attributes CAS = CAN001 Clip Clip CAS = CNA Settlement polyline layer Roads (ROAD) Select by attributes CAS = VIA Clip DEM Digit. Former river arms (FR) Zonal statistics as a table Elevation metrics tables Elevation metrics tables Elevation metrics tables Elevation metrics tables Elevation metrics tables … Delta River mouth (RM) Clip Near Coastal lagoon pres/abs map Tidal flat pres/abs map Salicornia-type pres/abs mapsSalt meadows pres/abs maps …Rice fields pres/abs maps RC LAG BAY IB OC FR RM ROAD CHANN RICE Distance to terrain variables tables Settlement polygon layer CAS = EDI001 Buildings (BUI) Clip Fig. 18. Cartographic model to assess the current distribution of wetland habitats
  • 42. 42   Generating data for the predictive model Once the dataset of habitat maps was developed, the next step was to create a random sample of 1.000 points for each wetland habitat type over the whole Delta plain to achieve data of the presence/absence of the habitats. The process was sequential again, and the steps are specified in the cartographic model (Figure 19). Several layers obtained in the previous steps (see database development) have been taken into consideration as excluding layers to overlap with the sample points. These geographic areas are: river channel, lagoons, irrigation channels, roads, buildings and rice fields. We excluded theses areas while the model was developed because they are parts of the delta where the habitats will be neither present and absent due to its land use. To sum up the process, the following steps are listed: a. Merging shapefiles of infrastructure elements (roads, channels and generic buildings) to obtain the constraining layer. b. Merging the infrastructure layer with hydrological layers (river channel, lagoons and rice fields). c. Creating a random point dataset (n = 1000) along the Delta as geographical limit layer. For each habitat type a set of layer random points was created (n = 9). d. Overlap data point layer with the excluding layer (infrastructures + water). The output layer was defined as points where the habitat is not present (layer absence points). e. For obtaining the layer of presence points of each habitat, a new random point dataset with the same number of absence points (to avoid bias in the sampling process) was created. The limit layer was the specific habitat presence map. f. Rasterization of vector layers for obtaining digital data of the independent variable over whole Deltaic plain: distances to rive/sea influence and anthropogenic limits. For this purpose the Euclidean distance tool of Spatial analysis extension was applied for each layer with 1m of pixel resolution. g. Each sample point layer (presence and absence) has its respective value of elevation, distances to river/sea influence and distances to hydrologic alteration.
  • 43. 43   h. The extraction of variables was executed with the Sample tool for obtaining the final data matrix with the 11 independent variables and presence/absence of each habitat type for total sample points.
  • 44. 44   Channel (CHAN) Roads (ROAD) River channel (RC) Lagoons (LAG) Buildings (BUI) Constraining layer Delta Create random points model_points n = 1000 absence_points n points = a absence_points absence_points absence_points absence_points Coastal lagoon pres map Tidal flat pres map Salicornia-type pres maps Salt meadows pres maps …Rice fields pres maps Create random points n = a absence_points absence_points absence_points absence_points presence_points Erase Rice fields (RICE) RC LAG BAY IB OC FR RM ROAD CHANN RICE RC LAG BAY IB OC FR RM ROAD CHANN RICE Z Coastal lagoon Tidal flat Salicornia-type Salt meadows Rice fields … Data matrix for each habitat type Sample Sample Euclidean distance 1x1m Fig. 19. Cartographic model to generate sample points and extract the metrics for predictive wetland model
  • 45. 45   3.7 Statistical analysis For each wetland habitat type, the presence/absence points and its respective terrain metrics (elevation, distances to river/sea influence and distances to hydrologic alteration sources) was determined with GIS-approach as described in the previous sections. PCA analysis was initially performed to study the overall relations between elevation and distance variables. Regression scores were extracted to analyze each wetland habitat pattern. Moreover, differences on variables between wetland habitats were assessed for achieving their ranges into the hydrological influence and soil altitude gradients. All the environmental independent variables were checked previously for normality and linearity. All the variables required natural logarithmic transformation to meet these parametric assumptions. Differences on vertical soil elevation between habitats assessed by CORINE land cover and mean transects was tested by Mann-Whitney test. Correlation analysis between elevation metrics (min, max, range and mean) and distances was performed to check the independence of the variables. All the statistical analyses were performed using SPSS v18.0 for Windows package. Logistic multiple regression predictive model As habitat map points are categorical variables (presence or absence), logistic regression is proposed for predicting occurrence of wetland habitat types from topographical deltaic variables (independent variables)(Hosmer and Lemeshow 2000). LMR is adequate because the dependent variable is dichotomous (presence/absence) and the model admits non- Gaussian independent variables (Franklin 1995). As explained, several authors proposed to use equal number of presence and absence points for developing logistic method to avoid bias (Felicísimo 2003; Narumalani et al. 1997). The foundation of the method is based on the probability of occurrence of any number of classes of a dependent variable (in this study, wetland habitat) based on explanatory variables (i.e elevation and distances). According to Custer and Eveleigh (1986), “regression modeling involves the derivation of a mathematical relationship between a set of independent predictor variables and a specific dependent condition”. A multiple logistic regression analysis within the sample points generated in previous steps resulted in an individual regression equation for each wetland habitat type.
  • 46. 46   Logistic regression is sensitive to extremely high correlations between variables that are supposed to be independent (Hosmer and Lemeshow 2000). To avoid this, high correlation between variables was eliminated by retaining only the variable with the highest explanatory power for pairs of variables with r Pearson coefficient (r > 0,6)(Filipe et al. 2002). In the multivariate analysis a forward stepwise was applied to each selected variable with a probability of entry of 0,05 and removal of 0,10. The addition and exclusion of variables was based on Wald’s test and the assessment of correlation was based on differences in the coefficients estimated when a variables is added to the model and from partial correlation of the estimated coefficients for p<0,001 (Zar 1999). To assess the fit of each model, the Chi-square test and a classification table was used. This procedure has been used by other authors (Narumalani et al. 1997; Ríos et al. 2005). The fit test examined the deviance of the model with the constant versus the final model and rejection was at p<0,05 significance based on a chi-square distribution. Another technique has been used to evaluate each predictive model. The Receiver Operating Characteristic (ROC) provides a threshold independent measure of accuracy and results in a plot of the relative proportions of correctly classified sites over the whole range of threshold levels (Narumalani et al. 1997; Tarkesh and Jetschke 2012). The ROC plot is obtained by plotting the sensitivity of the model against the false positive fraction over all thresholds, being the area under the curve (AUC) the probability that the model will distinguish correctly between observations. An area of 1 is perfectly accurate, whereas on of 0,5 is performing a random model (no acceptable). This fitting method has been applied in several studies of predictive vegetation modelling (Álvarez-Arbesú and Felicísimo 2002; Felicísimo et al. 2002; Syphard and Franklin 2010). All the statistical analyses were performed using SPSS v18.0 for Windows package. Model validation The predictive model has been validated with independent data obtained by field surveys, in which information about the presence/absence of different habitats was obtained by mean of transects. So, the validation was applied to Salicornia marshes, salt meadows and reed beds.
  • 47. 47   3.8 Model implementation in the GIS Logistic regression was used to generate probability models of habitat distribution with the introduction of a spatial component through GIS. Several authors have already applied it to coastal ecosystems (Álvarez-Arbesú and Felicísimo 2002; Narumalani et al. 1997; van Horssen et al. 1999; vandeRijt et al. 1996) and other fields, such as forested areas (Felicísimo et al. 2002; Turner et al. 2004) or soil mapping (Giasson et al. 2006). The LMR technique yields coefficients for each variable based on data derived from samples taken across a study site. These coefficients serve as weights in an algorithm which can be used in the GIS database to produce a map depicting the probability of wetland habitats. Quantitatively, the relationship between the "occurrence habitat" and its dependency on topographic variables can be expressed as: ( ))()(...)1()1()0( 1 1 )( nxnbxbb e iP ⋅++⋅+− + = where P(i) represent the probability value, x(1) … x(n) the values of the terrain variables and b(1) … b(n) are coefficients derived from logistic regression. Each regression equation results in a response value on an interval scale between 0 and 1. These response values can be interpreted as a relative frequency of occurrence or an estimate of probability of occurrences of wetland habitats. Then, low responses values will indicate low relative occurrence while high response values indicate high relative occurrence. The probability of occurrence was calculated from the logistic regression models in the raster calculator of ArcGIS. The coefficients of terrain variables which resulted statistical significant in each logistic model was applied to the GIS of Ebro Delta to produce a probability map of occurrence (cell size = 1m.) for two wetland habitats: Salicornia-type marshes and salt meadows.
  • 48. 48   4. Results and discussion 4.1 Current distribution of wetland habitats in the Ebro Delta Each wetland habitat was mapped and its extension was extracted in order to provide a baseline data for wetland cover in the study area. The analysis and elaboration of these maps through the existing CORINE land cover maps provided the more representative wetland covers in the Ebro Delta. A total of nine habitats were analyzed by an elevation dependent- approach, distance to river/influence and distance to sources of hydrologic alterations sources. Table 5 presents the general descriptors about habitat occupation in the delta: Table 5. Wetland habitat area in the Ebro Delta and its corresponding relative occupation. Urban areas and crops other than rice are excluded. *Priority habitat. Surface area of Delta = 330 km2 (Ibàñez et al. 2010).   Habitat EU code Patch number Area (km2) % Habitats % Delta Coastal lagoons *1150 34 16,85 5,86 5,11 Sandy habitats 2110, 2120, 2210 27 12,31 4,28 3,73 Tidal flats 1140 8 13,08 4,55 3,96 Salicornia-type marshes 1410 38 10,65 3,70 3,23 Salt meadows 1420 2 0,52 0,18 0,16 Cladium marshes *7210 7 3,35 1,17 1,02 Reed beds 53.1 29 8,67 3,01 2,63 Rice fields 82.d 10 222,05 77,17 67,29 Riparian vegetation 92A0 1 0,23 0,08 0,07 Total 287,75 Total surface occupied by wetland habitats was 287,75 km2 . Overall, rice fields are the dominant habitat in the deltaic plain in terms of % coverage of the study area. Sandy habitats (dunes and beaches) and tidal flats occupied near 10 % of the delta habitat surface, followed by salt marshes with Salicornia and Juncus (4%). Helophytic habitats, represented in the Delta principally by fresh and brackish marshes Cladium-type and reed beds, occupy 1,17 and 3,01% respectively.
  • 49. 49   Phragmites marshes (reed beds) have an important representation in the deltaic landscape since occupy brackish (Garxal), fresh marshes (Vilacoto) and the altered margins of the salt marshes due to agricultural runoff (Encanyissada, Aufacada). This habitat forms a great diversity of plants associations from a physiognomic point of view due to high variability of flooding levels and water salinity in the Delta (Curcó 2001). Under natural conditions (i.e peat soils largely flooded by carbonate fresh waters) Cladium- type marshes is limited to the natural wells “ullals” and some coastal lagoons that receives a significant of fresh groundwater supply (Encanyissada, Vilacoto). The potential area of Cladium marshes havs been modified by human activities (mainly agriculture) and its hydrological pattern has been altered by the establishment of an extensive draining system to lower the underground water level (Capítulo et al. 1994). This habitat is included in Annex 1 of the European Union Directive as a priority habitat type (7210 Calcareous fens with Cladium mariscus and species of the Caricion davallianae). The aquatic habitat of coastal lagoons is the other European priority habitat type included in our study. Coastal lagoons of the Ebro Delta are the most representative aquatic habitats and they occupy a higher surface area than the terrestrial habitats such as sand-dunes systems or Phragmites beds. Their hydrologic conditions have changed over the past 100 years due to the increment of the fresh water inputs and according Curcó and col. (1995) former coastal lagoons occupied most their potential area and were more numerous. There is a high variability in terms of surface of the lagoons: ranging from Garxal lagoon with 235 ha to Encanyissada lagoon with 786 ha. The main source of variation in the coastal lagoons is their hydrological regime which varies according to the fresh and salt water inputs. The freshest are the west basin of the Encanyissada lagoon, les Olles and those having more connection with the river (Garxal lagoon). The saltiest lagoons (la Tancada, Canal Vell or Buda), as other Mediterranean coastal lagoons, have a hydrological pattern linked with seawater fluctuations and some of them have periods of hypersalinity in summer (Badosa et al. 2006; Pérez-Ruzafa et al. 2005).
  • 50. 50   Within the elevation gradient, the area occupied by wetland habitats in the 0,0 – 0,3 m range is 73.54 km2 (26% of study area). From 0,3 to 0,5 m the surface occupied by wetlands is less (25.31 km2 , 9%). The habitats surface between 0,5 to 1 m have been 7,5 km2 (2.6% of study area). But between 1 and 1,5 meters of topographic elevation is where the largest area of habitats is concentrated due to the presence of major part of rice fields in this elevation range (168,22 km2 , 58%). The wetland area located beyond 1,5 meters only represents 0,31% of the study area, being the sand-dune systems the more representative habitats in this elevation range. The wetland habitats with mean elevations under mean sea level (-0,5 – 0,0 meters ) represent the 4,3 % of the total area (12,31 km2 ). In this range we find mainly coastal lagoons, tidal flats and reed beds. Table 6 summarizes the descriptive metrics of surface elevation for each habitat type included in the study: Table 6. Surface elevation metrics (min, max, rang and mean) for each wetland habitat. Values are shown in median and SE (in brackets); n= number of polygons. Habitat type n Min Max Range Mean Coastal lagoons 34 -0,325 (0,055) 1,718 (0,138) 2,044 (0,146) 0,181 (0,038) Sandy habitats 27 -0,145 (0,034) 2,731 (0,239) 2,875 (0,248) 0,698 (0,066) Tidal flats 8 -0,314 (0,072) 1,718 (0,227) 2,031 (0,259) 0,227 (0,089) Salicornia-type marshes 38 -0,205 (0,052) 2,204 (0,156) 2,409 (0,167) 0,509 (0,036) Salt meadows 2 0,410 (0,640) 2,275 (0,255) 1,865 (0,385) 1,125 (0,359) Cladium marshes 7 -0,302 (0,171) 1,692 (0,292) 1,994 (0,356) 0,387 (0,134) Reed beds 29 -0,690 (0,087) 1,900 (0,140) 2,590 (0,129) 0,301 (0,042) Rice fields 10 -0,623 (0,194) 3,625 (0,582) 4,249 (0,628) 0,820 (0,255) Riparian vegetation 1 0,490 4,230 3,740 2,808 Comparing the distribution of the eleven types of wetland habitats with the surface elevation of the Ebro Delta, we conclude that the existence of zonation is low but differences were found between habitats (Figure 20). One-way ANOVA tests (Table) indicate that there are
  • 51. 51   significant differences between the surface elevation of the wetland habitats observed. Elevation metrics (min, max and mean) were previously log-transformed for achieving parametric assumptions. A B C Reedbeds Ricefields Lagoons Tidalflats Cladium Salicornia Sand-dune Saltmeadows Riparian Reedbeds Ricefields Lagoons Tidalflats Cladium Salicornia Sand-dune Riparian Saltmeadows Cladium Reedbeds Tidalflats Lagoons Salicornia Salt meadows Ricefields Riparian Sand-dune -1,000 0,000 1,000 2,000 a b a,b a,b a,b a,b a,b a,b a a a a,b a,b a,b a,b b a a a,b a,b a,b b,c b,c c 1,000 2,000 3,000 4,000 5,000 0,000 0,500 1,000 1,500 2,000 2,500 3,000 Fig. 20. Surface elevation metrics (in meters) for each habitat type of the Ebro Delta. A: minimum, B: maximum, C: mean. The error bars represents the standard error considering all the polygons of the CORINE habitat map. Different letters point to significant differences (post hoc Tukey test: p < 0.05)  
  • 52. 52   Table 7. One-way ANOVA results for the surface elevation metrics extracted from CORINE land cover maps. Riparian vegetation was not included in the ANOVA analysis due to an insufficient number of samples. Elevation metrics Sum of Squares df Mean Square F Sig. Minimum Between Groups 2,219 8 0,277 5,652 <0,000 Within Groups 7,163 146 0,049 Total 9,382 154 Mean Between Groups 0,465 8 0,058 13,408 <0,000 Within Groups 0,637 147 0,004 Total 1,101 155 Maximum Between Groups 1,113 8 0,139 5,566 <0,000 Within Groups 3,676 147 0,025 Total 4,789 155 Range Between Groups 0,873 8 0,109 4,387 <0,000 Within Groups 3,655 147 ,025 Total 4,528 155 Lower elevations, excluding coastal lagoons, are occupied more frequency by tidal flats and reed bed habitats. Phragmites marshes are present in the lowest mean elevation in contact directly with coastal lagoons (Fig. 20A), as mapped in the CORINE land cover. Even though, the minimum elevation of emergent reed beds (-0,69 m.) has to taking into account since the growing of these communities is limited in waters of 0,3 – 0,4 m. depth (Coops et al. 1996; Squires and Valk 1992). Although scarce in the Delta (Ibàñez et al. 2002) this habitat type will be present in permanent or nearly flooded soils of fresh-brackish marshes. In salty coastal environments like the Ebro Delta, salt marshes are dominated by Salicornia- type vegetation and depending on its relative soil elevation, different genera can dominate (Ibàñez et al. 2010). In this study, Salicornia-type habitat had a mean elevation of 0,51 ± 0,04 m. Other authors (Pont et al. 2002) have found a similar topographic distribution of these habitats in the Rhône Delta, which range between 0,25 and 0,60 m. A more clear variation of soil elevation at upper regions was detected between Salicornia- type habitat and salt meadows dominated by Juncus. Even though this halophytic habitat has been drastically reduced in the Ebro Delta, its still occupies a broad range of elevation. Regarding salinity, salt meadows with Juncus maritimus and Juncus acutus occur in soils less
  • 53. 53   influenced by the underground sea water level in contrast to the exclusive halophytic communities (fruticose salt-marshes) (Espinar 2009). Thus, significant differences in mean soil elevation of Salicornia-type and Juncus-type environments were found (unpaired t test, p < 0,05). Silvestri and col. (2005) found a mean elevation difference between Juncus genus and Arthrocnemum genus of 15 cm. Our results show that given habitats are found at higher differentiated topographic elevations. The riparian habitat was observed at the highest mean and maximum surface elevation of the habitats (Fig. 20.D). The only patch mapped from CORINE land cover (Sapinya Island, 24 ha) shows a mean surface elevation of 2,81 m and its elevation can be considered representative of this habitat. This habitat is present in the fluvial levees that are the highest areas of the Delta plain, where it should develop according to the lowest salinity levels and eventual flooding events. Presently, the potential area of riparian vegetation of the Delta, especially Populus and Salix genus, has been transformed into rice fields and other crops. The humanized habitat of rice fields is present in the maximum range soil elevation extracted by the DEM of Deltaic plain. (Fig. 20.C). While other habitat types have a narrow elevation range (i.e tidal flats or Cladium marshes) rice fields exhibit a wide distribution, indicating a relative indifference to soil elevation. The high surface area occupied by rice throughout the Delta plain, from near-river lévées to the margins of the coastal lagoons has led to major part of the topographic gradient being occupied by this habitat. Moreover, the soil elevation of the Delta has been altered in many areas by agricultural purposes, lowering the upper zones and filling depressions. Then, the wide range in the elevation of this habitat could be attributed to this human factor. Methodological constrains on DEM application In several habitats, no consistent results have been detected in the application of high precise Digital Elevation Model of the Ebro Delta. Coastal lagoons showed mean elevation above sea level (0,18 m.), and its maximum elevation was placed around 1,7 m. When extracting elevation, the presence of micro-topography like “tores” (accumulation of soil in inundation areas that it elevates above water surface) probably result in a bias of elevation values. The level of detail in mean elevation of tidal flats (0,23 m.) it may associated to the same issue.
  • 54. 54   Depending on tidal range, this habitat should be located almost at sea level water or below sea level (e.g. -0,3 m.)(Sakamaki et al. 2006). In contrast, the spatial resolution of the DEMs in several studies where elevations are sampled at 30 m intervals, are more appropriate since its extension of the study area varies over thousands of square kilometres. (Brown 1994). Relationship between surface and soil elevation of wetland habitats The figure (Figure 21) shows the area occupied by natural habitat types (except rice fields) along the elevation gradient of deltaic plain. Habitats located in lowest elevations seem to be associated with marine-influenced environments, which coastal lagoons and tidal flats have maximum surface between 0,0 and 10 cm above mean sea level. Freshwater marsh habitats such as Phragmites-type occurs in high frequency on 0-1 and 0-2 m soil elevation directly on contact with water bodies that allow it flooded soils. Cladium marshes have showed a flat surface distribution as evidenced by its minimum topographic position more elevated than reed beds. Geographic position of Cladium habitats, near the inner border of the Delta associated with continental groundwater discharge areas, could explain its surface elevation pattern. Sandy habitats (dunes and beaches) are the habitat type with maximum surface at ca 0,5 m. Among this habitat, we can find several dune environments with different stability stages. This succession pattern (i.e embryonic, shifting and fixed) largely determines their topographic position along the elevation gradient of the Delta. Thus, according with the Delta prevailing wind (NW), we can find different patterns of sand-dune surface occupation due to the different orientation of the coast (Curcó 2006). At 0,6 – 0,7 m above the mean soil elevation, a few patches of Juncus with relative surface overlap with Salicornia-type marshes, especially in La Tancada. Finally, riparian vegetation has shown its maximum surface in the highest zone, although its representation in fluvial lévées has decreased considerably.
  • 55. 55   Area(km2)   Fig. 21. Distribution of natural habitat types as function of soil elevation and surface. Rice fields were not included in the plot.
  • 56. 56   Relationship between terrain variables For the entire wetland habitats the lowest elevation, highest elevation, range elevation and mean elevation show strong positive correlation (Pearson, p<0,01). The mean elevation has been correlated positively with the distance to channels and negatively with distance to inner border, lagoons, river and former river arms. The three variables related with hydrologic alterations elements (roads, channels and rice fields) were strongly (negatively) correlated to each other, which was expected due to their overlapped position in the deltaic plain. Note that distance to former river arms of the Ebro River (Riet de Zaida, Fondo and Muntells) was correlated positively with distance to rice fields, channels and roads of the Delta. This can be explained because of these ancient courses are largely occupied by rice fields and consequently by channels, either irrigation or drainage. A significative (negative) correlation was found between distance to bay and river channel that could represent a longitudinal gradient from fluvial lévées to bay (marine influence). Distances associated with the riverine influence are also strongly correlated, mainly between river channel and river mouth. PCA ordination In order to investigate the relations between all the environment variables on wetland habitats distribution a principal component analysis (PCA) with varimax rotation was carried out (Figure 22). Most of the analyzed variables were interdependent and have significative correlation among them (Table correlations). The usefulness o the PCA was checked through Kaiser-Meyer-Olkin’s (KMO) measure of adequacy sample (0,686) and Bartlett’s test of sphericity (p<0,001). The two first axes explaining the 26% and 22% of the total variation respectively. The minimum elevation of habitat polygons and distances to anthropogenic elements was correlated positively with PCA axis 1. The first axis separated the patches of wetland habitats closer to road, channels rice fields and former river arms with higher minimum elevation than patches placed far of these elements (lower minimum elevations). Then, the first axis summarizes the variation associated with rising elevation of wetland habitat placed near anthropogenic elements and ancient river lévées. PCA axis 2 explains the variation associated with maximum elevation gradient of habitat patches from interior to exterior deltaic plain. The highest elevation of habitats patches has been correlated
  • 57. 57   (positively) with areas placed near the inner border of the Delta and opposites to exterior limits of Delta (outer coast and river mouth). The regression scores of each habitat polygon were extracted to visualize their position on the two PCA axes. Analysis of salt meadows and riparian vegetation were not assessed due to its lower polygons in the CORINE habitat map. Some coastal lagoons were relatively separated on PCA axis 2: Buda lagoons were negatively correlated with it (i.e. near from river mouth and outer coast and lower maximum elevations) and patches of Encanyissada were correlated positively (i.e. far from river mouth and higher maximum elevations). Sand dunes showed patches separated along PC2. Sandy habitats correlated positively with this axis have higher maximum elevations and are influenced by river mouth and outer coast (e.g. Fangar dunes). While dunes/beaches oppositely placed along PCA axis 2 have shown lower maximum elevations and were more influenced by bays and inner border (e.g. dunes/beaches of Trabucador barrier). Tidal flats polygons have been differentiated in the patches of Buda area (near river mouth and correlated positively with PC1) and patches of Punta de la Banya. The patches of Salicornia habitats were relatively separated along PCA axis 2. Then, Salicornia-type marshes of Buda were different of polygons placed in la Banya given that the last showed negative correlation with elevation and distance to river mouth. It means that while there are some patches of Salicornia with lower maximum elevations located near the bay, oppositely other groups with higher maximum elevations were placed near river mouth. The reed beds were grouped according to PCA axis 2 mainly. Then, polygons with lower maximum elevations and near to river mouth and outer coast were correlated negatively with PC2 (i.e. reed beds of Buda lagoon as representative place). While patches of Phragmites located in la Tancada seems to be associated with higher maximum elevations and greater distances to these limits than either Buda or Garxal reed beds (positively correlated with PC2). PCA axis 1 explains another source of variation related with reed beds distribution. This axis separates groups of reed beds placed far from anthropogenic and former river arms which can cause uprising elevation effect (higher minimum elevations).
  • 58. 58   The ordination technique has been useful to assess the relative distribution of wetland habitats according to maximum variation sources of topographic deltaic variables. However, the forcing factors of wetland distribution include other variables as soil salinity or moisture content (Moffett et al. 2010). Regarding elevation, we can expect that habitats with higher vertical elevation will have well-drained soils and lower flooding periods. Linking with the soil salinity, the duration of evaporation periods (occurring when the marsh is not flooded) increases with elevation and thus salts become increasingly concentrated (Adam 1993). Then, stress conditions associated with salinity will be more evidence in lower regions within intertidal range of the Ebro Delta (0-0,5 m)(Jiménez 1996). Soil salinity decreases beyond sea water influence, therefore, these observations indirectly concerning the presence of wetland habitats to topographic position of Delta.
  • 59. 59     Fig. 22. PCA-Ordination diagram of the environmental variables included in the study. For abbreviations see the methods section.
  • 60. 60   Table 8. Mean soil elevation, distances to river/sea influence and distances to hydrological alteration of the main habitats in the Ebro Delta. Standard error of mean in italics. Habitat Mean elevation (m) Dist. to outer coast (km) Dist. to inner border (km) Dist. to river channel (km) Dist. to lagoons (km) Dist. to bay (km) Dist. to river mouth (km) Dist. to former river arms (km) Dist. to rice fields (km) Dist. to channels (km) Dist. to road (km) Coastal lagoons 0,181 1,549 11,381 6,409 0,000 3,933 13,078 6,278 1,517 0,141 0,085 0,038 0,344 0,801 0,953 0,000 0,607 1,511 0,743 0,456 0,048 0,031 Sandy habitats 0,698 0,656 8,325 9,265 1,046 1,608 16,406 7,401 2,825 0,977 0,758 0,066 0,160 0,842 1,073 0,210 0,441 1,905 1,136 0,503 0,246 0,244 Tidal flats 0,227 0,718 7,328 12,677 0,663 1,658 23,103 12,368 4,178 1,523 1,493 0,089 0,183 1,572 1,578 0,351 1,248 2,922 0,953 0,629 0,722 0,724 Salicornia-type 0,509 0,817 9,955 9,801 0,909 2,431 17,780 8,561 3,105 0,677 0,585 0,036 0,181 0,674 0,979 0,173 0,432 1,540 0,911 0,492 0,165 0,162 Salt meadows 1,125 4,559 7,301 4,237 0,502 2,916 18,096 4,489 0,000 0,000 0,000 0,359 4,342 6,829 0,231 0,135 0,884 6,676 3,851 0,000 0,000 0,000 Cladium-type 0,387 7,632 1,891 6,257 0,060 2,092 22,352 7,359 0,063 0,006 0,085 0,134 0,467 0,382 0,629 0,044 0,542 0,570 0,514 0,041 0,004 0,031 Reed bed 0,301 1,582 11,944 3,207 0,152 4,418 7,071 3,140 0,086 0,039 0,058
  • 61. 61     Table 8 continued 0,042 0,278 0,855 0,426 0,060 0,563 0,905 0,453 0,049 0,020 0,037 Rice fields 0,820 2,695 6,849 1,854 0,601 3,740 9,660 2,734 0,000 0,000 0,000 0,255 1,287 2,105 0,926 0,598 0,949 2,450 0,962 0,000 0,000 0,000 Riparian vegetation 2,808 10,537 3,674 0,000 6,215 8,458 18,067 1,686 0,201 0,166 0,055 0,000 0,000 0,000 0,000 0,000 0,000 0,000 0,000 0,000 0,000 0,000 ANOVA test p<0,0001 p<0,0001 p<0,0001 p<0,0001 p<0,0001 p<0,0001 p = 0,215 p<0,0001 p<0,0001 p<0,0001 p<0,0001
  • 62. 62   Table 9. Pearson’s correlation coefficients among independent variables for the current distribution of the habitats type in the Ebro Delta. Significance levels ** p<0,01; * p<0,05. mean ELEV min ELEV max ELEV range ELEV OC IB RC LAG BAY FR RM RICE CHANNEL ROAD mean ELE 1 min ELE ,396** 1 max ELE ,618** ,110 1 range ELE ,374** -,283** ,899** 1 OC -,067 -,054 -,287** -,259** 1 IB -,323** -,047 -,332** -,288** -,143 1 RC -,218** ,012 -,443** -,423** ,233** ,014 1 LAG ,408** ,323** ,014 -,151 ,114 -,057 ,220** 1 BAY -,024 ,008 -,172* -,189* -,096 ,385** -,275** -,179* 1 FR -,198* ,215** -,212** -,314** -,002 ,425** ,247** -,021 ,105 1 RM ,017 ,027 -,280** -,298** ,305** -,196* ,698** ,281** -,257** ,063 1 RICE ,050 ,423** -,090 -,281** ,060 ,068 ,240** ,366** -,188* ,474** ,135 1 CHANNEL ,184* ,416** -,006 -,206* -,001 ,036 ,211** ,298** -,089 ,418** ,144 ,749** 1 ROAD ,030 ,359** -,050 -,203* ,034 ,038 ,129 ,137 ,141 ,379** -,003 ,605** ,703** 1 OC: Distance to outer coast IB: Distance to inner border RC: Distance to river channel LAG: Distance to lagoons BAY: Distance to bays FR: Distance to former river arms RM: Distance to river mouth RICE: Distance to rice fields CHANNEL: Distance to channels ROAD: Distance to roads
  • 63. 63   Distances to riverine/marine and human infrastructures Mean distances to river/sea influence (Figure 23) and human infraestructures (Figure 24) for each wetland habitat are plotted (Table 8). In general, the hydrological boundaries associated with marine influence (the outer coast and bay mainly) have the lowest distances to wetland habitats like tidal flats, sandy habitats (dunes and beaches) and Salicornia-type marshes. Other habitats, such as Cladium-type marshes or riparian vegetation showed higher distances to those variables (post hoc Tukey test p<0,05). Phragmites marshes and salt meadows instead were located in intermediate distance within this marine influence gradient. The effect of permanent flooded areas (i.e coastal lagoons) has been demonstrated with the presence of emergent helophytic vegetation (reed beds and Cladium marhes) placed closer than Juncus meadows. However, the few samples cases of this habitat type (salt meadows) didn’t allow to assess the influence of hydrological boundaries in a clear form. Except riparian vegetation, tidal flats, salt marshes, sandy habitats and rice fields forms an homogenous group in relation with distance to lagoons (Post hoc Tukey, p<0,05). That is, there are no statistical differences in the distance to coastal lagoons of these wetlands habitats. These results confirm the position of several habitats like patchwork of different classes without a clear pattern around the lagoons. Riverine influence The geographic position of habitats according to its distance to inner border shows no clear pattern, even though coastal habitats (i.e dunes and beaches, tidal flats), as expected, were found at higher distances. The riverine influence expressed as distance to river channel and river mouth mainly shows that riparian vegetation, reed beds, and rice fields, in this order, are the closer habitats to these limits. The ANOVA tests showed no significant differences in distance to river mouth of the wetland habitats (p = 0,215). A post hoc Tukey test indicated that the differences in mean distance to river channel increases from reed beds and rice fields to all the other habitats. The same results were found when the position of habitats respect to former river arms is considered. In figure 23, we can observe increase in distance to riverine influences from “fresher” habitats (closer) to marine habitats (far away). That is, tidal flats, Salicornia
  • 64. 64   marshes and sandy habitats were placed far away from river channel, river mouth and former river arms. Human infrastructures influence Distances to hydrological alteration sources have shown significant differences of wetland habitats studied (ANOVA test, p < 0,001 in all cases) (Figure 24). Reed beds, salt meadows and Cladium marshes were the habitats closer to rice fields and channels (mean distance < 100 m.) while Salicornia-type marshes, tidal flats and sandy habitats were located far away (Post hoc Tukey test p<0,05). The close position of rice fields, channels and roads to coastal lagoons (mean distance = 0,58 km), in contrast to other habitats, such as Salicornia salt marshes (mean distance = 1,45 km) makes evidence supporting the hypothesis of a hydrological alteration of closest alteration elements to lagoons . The proximity of the disturbing elements can produce different effects depending on their potential effects: hydrology balance between fresh (agricultural runoff) and salt water, and barrier effect. Thus, effects of rice fields will base on the variation of hydrologic balance between fresh water inputs during irrigation period and salt water conditions during the rest of the year. Consequently, aquatic vegetation of coastal lagoons is undergoing changes in their communities (Menéndez and Comin 2000; Menéndez et al. 2002). On the other hand, roads and tracks may act as barriers of hydrological fluxes, and therefore, natural flows of water have been altered. The effects of the proximity of channels on wetland distribution are difficult to discern using the approach of the present study. Setting aside different hydrological effects (i.e drainage or irrigation water), we can assume a barrier effect of these elements in the water natural flow. So, the fact that the habitats are closer to hydrological alteration elements does not mean a greater effect on their distribution.
  • 65. 65   0 2 4 6 8 10 12 Sandy habitats Tidal flats Salicornia-type Coastal lagoons Reed bed Rice fields Salt meadows Cladium-type Riparian vegetation Distance to outer coast (km) 0 2 4 6 8 10 12 14 16 Rice fields Riparian vegetation Cladium-type Reed bed Salicornia-type Tidal flats Coastal lagoons Sandy habitats Salt meadows Distance to inner border (km)   0 1 2 3 4 5 6 7 8 9 Sandy habitats Tidal flats Cladium-type Salicornia-type Salt meadows Rice fields Coastal lagoons Reed bed Riparian vegetation Distance to bay (km)   0 1 2 3 4 5 6 7 Coastal lagoons Cladium-type Reed bed Salt meadows Rice fields Tidal flats Salicornia-type Sandy habitats Riparian vegetation Distance to lagoons (km)  
  • 66. 66   0 2 4 6 8 10 12 14 16 Riparian vegetation Rice fields Reed bed Salt meadows Cladium-type Coastal lagoons Sandy habitats Salicornia-type Tidal flats Distance to river (km)   0 5 10 15 20 25 30 Reed bed Rice fields Coastal lagoons Sandy habitats Salicornia-type Riparian vegetation Salt meadows Cladium-type Tidal flats Distance to river mouth (km)   0 2 4 6 8 10 12 14 Riparian vegetation Rice fields Reed bed Salt meadows Coastal lagoons Cladium-type Sandy habitats Salicornia-type Tidal flats Distance to former river arms (km)   Fig. 23. Mean distances (in km) to river/sea influence for wetland habitats of the Ebro Delta. The errors bars represent the standard error of mean.
  • 67. 67   0 1 2 3 4 5 6 Salt meadows Rice fields Cladium-type Reed bed Riparian vegetation Coastal lagoons Sandy habitats Salicornia-type Tidal flats Distance to rice fields (km)   0 1 1 2 2 3 Salt meadows Rice fields Cladium-type Reed bed Coastal lagoons Riparian vegetation Salicornia-type Sandy habitats Tidal flats Distance to channels (km)   0 1 1 2 2 3 Salt meadows Rice fields Riparian vegetation Reed bed Coastal lagoons Cladium-type Salicornia-type Sandy habitats Tidal flats Distance to roads (km)   Fig. 24. Mean distances (in km) to hydrological alterations for wetland habitats of the Ebro Delta. The errors bars represent the standard error of mean.