Proyecto Graze

Convocatorias 2016
Proyectos RETOS
Dirección General de Investigación Científica y Técnica
Subdirección General de Proyectos de Investigación

TÍTULO DEL PROYECTO: Desarrollo de sistemas sostenibles de producción ganadera en espacios protegidos con alta variabilidad interanual en la producción primaria: vacas, caballos y ciervos en el E.N. Doñana
En áreas mediterráneas y semiáridas, la elevada incertidumbre en el régimen de precipitaciones y la productividad vegetal asociada represent un importante reto para el desarrollo y optimización de sistemas sostenibles de producción ganadera extensiva, particularmente en áreas con alta abundancia de ungulados silvestres.  Este proyecto está enfocado endesarrollar criterios a largo plazo y herramientas de modelado para la gestión sostenible de ganadería extensiva en áreas protegidas mediterráneas con alta variabilidad climática, centrándose en el Espacio Natural Doñana (END) como caso de estudio. Para ello, propone desarrollar un programa interdisciplinar que combina: (1) La obtención de estimas históricas y actuales del tamaño poblacional, uso del espacio y requerimientos alimenticios de los cinco ungulados presentes en el END (silvestres: gamos, ciervos y jabalíes; domésticos: vacas y caballos). (2) Estimas históricas y actuales del efecto del nivel de precipotación y la presión de herbivoría sobre la producción de biomasa aérea de la vegetación de Doñana, basadas en una combinación de prospecciones de campo, experimentos y modelos de teledetección. (3) El desarrollo de modelos dinámicos y espacialmente explícitos de la relación entre precipitación, presión de herbivoría y producción de biomasa vegetal en el END, y su utilización para generar y evaluar diferentes escenarios de gestión en las condidiones climáticas actuales y aquellas previstas en los diferentes escenarios de cambio climático. El resultado esperado es el desarrollo de una base de conocimiento, de criterios a largo plazo y de herramientas de modelado para la gestión sostenible de la ganadería extensiva en Doñana y en otras áreas protegidas de la región mediterránea.



Free-ranging livestock production is a widespread exploitation system in many areas of the world (Launchbaugh & Howery 2005), including the Iberian Peninsula. The high conservation of many of these areas has resulted in their inclusion within conservation areas, where human exploitation systems are then requested to meet the societal demand of nature conservation. In many of these areas, often as a result of the management changes associated to its new status (e.g. hunting reduction or elimination, access restrictions to the local population), two conflicting trends arise: livestock must coexist with increasingly large numbers of wild ungulates, and local stakeholders may increase their demand for higher livestock stocking rates for a diversity of reasons (e.g. increasing economic gain, compensating the loss of other sources of income and/or gaining access to the conservation site) (Weisberg et al. 2002). The resulting management conflict often result in local or widespread overgrazing, reducing plant productivity and thus affecting both livestock production and nature conservation (Vavra et al. 1999) – and may develop into open social conflicts (Gómez-Baggethun et al. 2013, Bejarano-Bella & Torres Rodríguez 2016). Unfortunately, these effects often reflect a lack of adequate knowledge or understanding of the factors driving to effects of livestock on biodiversity (Lunt et al. 2007).

In strongly seasonal areas, such as the Mediterranean region, this problem may be exacerbated by the perceptual and management constraints imposed by the large seasonal and inter-annual fluctuations in rainfall, which results in large changes in plant productivity across years (e.g. Lázaro et al. 2001). In such situations, planning the stocking rates and management regime becomes increasingly harder owing to the amplification of effects of environmental unpredictability by the strong feedbacks between plant standing crop and regrowth potential (which amplifies the impact of grazing on plant growth) (e.g. Naveh 1982). In dry years, overgrazing exhausts the asexual organs and depletes the seed bank of the vegetation, thus reducing its productivity during the wet years that follow. And, while wild ungulates may even have positive effects of livestock during wet years (Odadi et al. 2011), the increases in population numbers of wild ungulates and/or in the stocking rates of livestock following (sequences of) wet years may also increase the likelihood of overgrazing in the dry years that follow. In all these cases, food limitation during periods of low productivity requires the assumption of additional expenses by livestock breeders (who must stock food to free-ranging animals or transport them to pens during such period) and/or may entail levels of ecosystem degradation that may be unacceptable in conservation areas.

The solution to this dilemma is complex, since it involves the use of very conservative stocking rates (low enough to prevent overgrazing in dry years) or to develop systems allowing for a dynamic adjustment of stocking rates and/or space use by livestock in response to rainfall levels (Graham et al. 2010, Weisberg et al. 2002). In such case, early detection of expected plant production levels based on combined records of rainfall levels and cumulative grazing pressure may represent a highly valuable tool, allowing for the adjustment of grazing rates before reaching levels conductive to the long-term degradation of plant production (Rowntree et al. 2004). The development of such tools is all the more valuable because climate change is expected to result in an amplification of the hydrological cycle, with more extreme intra-annual precipitation regimes characterized by larger rainfall events and longer intervals between events (e.g. Knapp et al. 2008) – thus resulting in an amplification of the processes described above.

The processes described above may however be buffered by landscape heterogeneity, particular when it is related to water storage and availability (due e.g. to changes in topography and soil types). In such cases, lowland areas may provide areas with high plant productivity in dry years; and these areas will be more resilient to overgrazing because they not accessible to herbivores (due to prolonged flooding) in wet years. This mechanism, which fits our preliminary observations in the Doñana marshes (the study system proposed for this project, see below) and was reflected traditional management method in the area (Gómez-Baggethun et al. 2012), may provide additional opportunities for the sustainable management of free-ranging livestock in conservation areas where wild ungulates are also present, through the wise adjustment of the stocking rates and space use by such herbivores to the rainfall levels of the current and previous years (Gómez-Baggethun et al. 2013). 

Study system
The project will take place at the Doñana Protected Area (Espacio Natural Doñana, END hereafter), a nature reserve (including a National Park and several areas of Natural park) located on the Atlantic coast of South-West Spain. The region has a Mediterranean climate, classified as dry sub-humid with marked seasons. Grazing by domestic and wild ungulates varies along the different seasons, following changes in plant productivity and accessibility at the different landscape units (Barasona et al. 2014). In the wet season (December–May), marshlands are flooded and ungulates graze in the sandy shrublands. As the dry season progresses and shrubland vegetation senesces, ungulates start grazing in the marshland (where either bulrushes, notably of Scirpus maritimus, or saltworts, gen. Sarcocornia, Arthrocnemum and Suaeda, tend to be the dominant vegetation). Ungulates become most food limited during summer (June–September), when wetlands dry up causing senescence of their herbaceous vegetation. The humid ecotone between the shrubland and the marshland (locally known as “vera”) remains humid throughout the year. Therefore its vegetation, dominated by herbaceous meadows (Galiopalustris sp. with Juncus maritimus associations), represent a key food supply for ungulates during such season.
The study area has moderate to high densities of red deer, fallow deer and wild boar. A traditional breed of cattle (locally called “marismeña”) is farmed within five cattle management areas in the END: Coto del Rey (CR), the northernmost area, contains no cattle husbandry; three areas in the center: SO (n = 350 cattle; density = 5.7 cattle/km2), BR (n = 168 cattle; density = 2.6 cattle/km2), and PU (n = 152 cattle; density =4.0 cattle/km2); and Marismillas (MA), the southern-most area (n = 318 cattle; density = 3.1 cattle/km2) (Barasona et al. 2014). Each cattle management area is surrounded by a cattle-proof fence, which limits the movements of each herd to within their designated management area. However, social groups (overwhelmingly females) showing individual ranging behaviour may be differentiated within each cattle management area (Lazo 1995).
Previous results of the research team and other national and international teams
This project builds on the synergies of the different research interests of the members of the work team and, while providing a step further in their own research lines, creates the opportunity of achieving an enriching synthesis of their present and previous work. The project aims indeed at combining the expertise of all these researchers in a collaborative and truly inter-disciplinary effort, with the purpose of addressing the interweaving factors and mechanisms that make-up the complexity of plant-herbivore interactions in a coherent and comparative manner, rather than dissecting and addressing them using piecemeal approaches. At the same time, it aims at providing knowledge and specific tools of direct practical use for the sustainable management of domestic livestock and wild ungulates in Mediterranean ecosystems, typically subjected to high inter-annual variation in key determinants of plant productivity (most notably, rainfall), which should be directly applicable to the case study (the Doñana Protected Area) and hopefully easy to extrapolate to other areas of similar climate and conditions.
Luis SANTAMARIA, a tenured researcher (Científico Titular) who leads the Spatial Ecology Group at the Doñana Biological Station (EBD-CSIC), has a extensive experience in the main topics addressed in this study, including (i) the ecology, evolution and management of plants and herbivores (e.g. Jonzen et al. 2002, Latorre et al. 2013, Santamaría 2002, Santamaría & Rodríguez-Gironés 2002, Santamaría et al. 2007), (ii) the ecology of the Doñana marshes and its vegetation (Santamaría et al. 1996, Santamaría et al. 2005, Santamaría & Amézaga 1999), (iii) the development of spatially-explicit models and its application to the management of conservation areas (e.g. Magrach et al. 2011, Rodríguez-Pérez et al. 2012), and (iv) the collaborative development of management plans and policies (e.g. Méndez et al. 2012, Santamaría & Méndez 2012, Santamaría et al. 2013). During such work, we successfully used all the techniques proposed in this project, such as surveying animal populations, tracking and modelling animal movement, surveting the composition and biomass yield of plant populations, applying remote-sensing techniques to habitat characterization and modelling, developing spatially-explicit models, and conducting collaborative workshops to
Luis Santamaría’s expertise will be complemented by the extensive knowledge of the work team, which includesRicardo DÍAZ-DELGADO, a tenured technician at EBD-CSIC who has an extensive experience in remote-sensing and GIS analysis of Doñana’s landscape and vegetation structure, as well as in the development of long-term monitoring programs (he is the Spanish representative at the International Long-Term Ecological Research network, ILTER). Francisco CARRO, a tenured technician at EBD-CSIC who has an extensive experience in monitoring wildlife mammals, including ungulates, at the Doñana region (he is the coordinator of the mammal monitoring program at EBD). Cristina PÉREZ, who has ample experience in the analysis of vegetation and herbivore-dropping samples.
Plant-ungulate interactions and, more broadly, plant and ungulate ecology are very wide disciplines with many different teams working on very different model organisms around the world. At international level, a few examples of leading research teams include those led by Marcelo Sternberg (Tel Aviv University), Peter Weisberg (University of Nevada), Mark Hebblewhite (University of Montana) and Philippe Ballon (IRSTEA, Antony, France), as well as interdisciplinary groups such as the Ungulate Ecology Group (CEES Norway). At national level, some of the leading research teams include those led by Juan Carranza (Universidad de Córdoba) and J. Vicente (IREC).
Purpose and opportunity of the Project

Finalidad del proyecto, oportunidad de llevarlo a cabo y adecuación del mismo a la Estrategia Española de Ciencia y Tecnología y de Innovación y, en su caso, a Horizonte 2020 o a cualquier otra estrategia nacional o internacional de I+D+i.

The develoment of sustainable livestock production systems is one of the key elements of Challenge 2 of the Spanish Strategy of Science and Technology (Reto 2: Seguridad y calidad alimentaria; actividad agrarian productive y sostenible, sostenibilidad de los recursos naturales, investigación marina y marítima), which explicitly mentiones the need to improve the competitiveness of the food-production sectors while improving the management of natural resources and  (“… la competitividad de los sectores agroalimentario, forestal y pesquero en los mercados nacionales e internacionales, …, mejorando la gestión de los recursos naturales utiliz ados por los distintos sectores productivos… [y] avanzar en la conservación de los recursos naturales, en particular en el uso eficiente del agua, … la lucha contra la erosión de los suelos, …, la protección de nuestros sistemas agroecológicos, su biodiversidad…”). More specifically, the Priorities set with Challenge 2 include explicitly the sustainable improvement of livestock production systems, including the improvement of its productive efficiency, the reduction and improvement of the intakes and the environmental and social valuation and modeling of agro-forestal systems (Prioridad II. Mejora sostenible de los sistemas de agrícolas, ganaderos y forestales: (i) eficiencia productiva … en especies agrícolas, ganaderas y forestales, promoviendo … el desarrollo y mejora de la eficiencia de los insumos; […] (iv) sistemas de producción animal y vegetal, incluyendo insumos, maquinaria, tecnologías y sistemas; […] (vi) valoración y modelización económica, ambiental y social de los sistemas agroforestales.”)

Similarly, the economic and environmental importance of sustainable food production systems, including the sustainable production of terrestrial livestock, is explicitly mentioned in Horizon 2020 Societal Challenge 2 on “Food security, sustainable agriculture, marine and maritime research and the bioeconomy”. Indeed, previous H2020 calls within this program have topics such as “Sustainable terrestrial livestock production”, framed within the more general topic “Sustainable food production systems”. Several parts of these documents and calls have stressed the importance of improving sustainability and productivity of terrestrial livestock systems, particularly in the face of global change, which is likely to increase the environmental footprint of such system. This project fits perfectly within such goal, by aiming to provide tools and criteria for the development of sustainable livestock production in systems where free ranging cattle and horses feeding on natural vegetation and share such food resources with wild ungulates – in particularly, at natural protected areas where such production must be harmonized with the need to protect highly-valued biodiversity and ecosystem services. Its focus, aim and interdisciplinary approach will contribute to meet the main objectives of the Programme, such as “test, demonstrate and transfer effective solutions to major challenges affecting the Bioeconomy on land and sea, across the agri-food chain from soil to society”,  “unlock the potentials of available bio-resources in the different bioeconomy and blue-economy sectors in a sustainable and socially responsible way”, and “bringing Research and Innovation at the heart of major primary sectors … taking advantage of new potential in the biological, ecological, technical and information technology domains”.

This project builds on the hypothesis that in Mediterranean and semiarid areas, large uncertainty in rainfall levels and the associated levels of plant primary production represents a strong contstrain for the optimal, sustainable and environmentally-responsible management of free-ranging livestock production, particularly for conservation areas such as the Doñana National Park. Overgrazing during dry years exhaust the asexual organs and depletes the seed bank of the vegetation, thus reducing its productivity during wet years; while the increases in population numbers of wild ungulates and/or stocking rates of livestock following (sequences of) wet years increases the likelihood of overgrazing when they are followed by dry years.

As a secondary hypothesis, we will evaluate whether landscape heterogeneity related to water storage and availability (due e.g. to changes in topography and soil types) may mitigate these effects by providing areas with high plant productivity in dry years, which are not accessible to herbivores (due to prolonged flooding) in wet years. This hypothesis fits our preliminary observations in the Doñana marshes.

Main Goal

The project aims at developing long-term criteria and modelling tools for the management of free-ranging livestock in Mediterranean protected areas with high climatic variability, taking the Doñana Protected Area (Espacio Natural Doñana, END) as a case study.

Specific Goals

1.       Estimate the population size and feeding demands of the five ungulates feeding in Doñana (wild: red deer, fallow deer, wild boar; domestic: cattle, horse).
2.       Estimate the effects of rainfall level and grazing pressure on the productivity and standing crop of the vegetation of Doñana.
3.       Reconstruct the long-term changes in productivity and biomass yield of the Doñana vegetation, using remote-sensing tools.
4.       Develop models of the relationship between rainfall, grazing pressure and plant standing crop at the END, and use them to evaluate different management scenarios under current and forecasted climate conditions.

Infrastructure y equipment available for the project

The centers and laboratories to which the members of the research team belong are equipped with all the equipment and material necessary for the development of the project (with the exception of the items included in the budget). The EBD-CSIC counts with basic lab equipment such as precision scales, digital calipers, fridges and freezers, stereoscopes and (visual and epifluoresce) microscopes, binoculars and telescopes, manual and digital refractometers, together with climatic and germination chambers, greenhouse facilities and animal experimentation facilities. EBD’s Service Units include fully-equipped Animal Physiology Lab, Molecular Ecology Lab and Aquatic Ecology Lab.

Use of ICTS-Doñana
The Doñana Scientific Reserve (RBD) is a Singular Research  Infrastructure (ICTS)
belonging to CSIC, which depends administratively from ERBD-CSIC. It extends over
10,000 ha within the 50,000 ha of the Doñana National Park and offers a complete infraestructure for research purposes. Around almost a third of our project will be developed in RBD. While we cannot apply for access to ICTS-RBD because we are staff of EBD (our travel and living costs will be funded from the current proposal), we will benefit from the use of its facilities during the extensive field work planned in Doñana. During the field campaigns there, the researchers have access to the field Wi-Fi network (allowing them to connect their devices, e.g. data-loggers), field transportation (4x4 vehicles), use of field laboratories and housing.


Ob.1. Estimate the population size and feeding demands of the five main herbivores feeding in the Doñana marshes (wild: red deer, fallow deer, wild boar, hare and rabbit; domestic: cattle, horse, sheep)

Task 1.1. Historical data on herbivore abundance and population structure
- Achievement H1: Assessment of historical changes in ungulate abundance completed (data collection and statistical tests).
- Deliverable E1: Scientific Article (together with Task 1.2. and 1.3.).
The population size (number of individuals) and population characteristics (sex, age class) of the six main herbivores present at the END have been registered in a number of management, monitoring and research surveys since the early 80’s. To our knowledge, however, they have not been gathered systematically and subjected to comparative analysis. In this WP we will gather all the available information from published and unpublished, scientific and technical sources (e.g. National Park and END management plans; RBD-ICTS monitoring program; Soriguer 1981, 1983, 1988; Lazo et al. 1991; Delibes &Soriguer 1993; Soriguer et al. 1994, 2001) and use them to reconstruct the long-term changes in their abundance.

Inasmuch as the available data allow for it, we will also develop simple demographic models to evaluate the effect of historical changes on resource availability (estimated using remote sensing, see below), inter-specific competition (within the herbivore’s guild) and management measures (e.g. culling of fallow deer, reductions and increases in the stock of domestic herbivores) on the survival, recruitment and population size of the different species.

Task 1.2. Spatial distribution and individual movements
- Achievement H2: Assessment of spatial variation in ungulate abundance, in relation to the availability and exploitation of food resources, completed (data collection and statistical tests).
- Deliverable E1: Scientific Article (together with Task 1.1. and 1.3.).

Previous studies have shown that the distribution of herbivores is not homogeneous across the Doñana Nature Reserve (END). The abundance of cattle and horse varies largely among the different management areas, which are separated by fences for the purpose of livestock management (see above). Wild ungulates, though generally able to cross such fences, also show large differences in abundance across states. As these studies acknowledge, the extent to which these differences (which are based on one to three censuses a year, plus trimestral surveys of droppings for a certain states and years) adequately reflect seasonal and diel changes in space and habitat use remains unknown. Some of these changes are strong determinants of the spatial gradients in grazing pressure, e.g. in the marsh area, where grazing progresses from shallower to deeper areas as the inundation recedes along the spring. Such knowledge is therefore essential to evaluate the extent to which herbivores compete for their food sources and/or they have synergic or merely additive impacts on their food plants.

To evaluate the importance of temporal and spatial variation in the local abundance (space use) by the different herbivore species, we will work at four different spatio-temporal scales: 

1.2.1. We will analyze the data available from the monitoring of four species of wild ungulates (red deer, fallow deer and wild boar) by the RBD-ICTS. These data consist of direct counts on 8 transects of approx. 15 km, recorded three times a year since 2005, and provide density estimates (#individuals/km) based on the adjustment of detectability curves (distance sampling).

1.2.2. We will complement these observations with the analysis of yearly counts of droppings (fecal items) at 24 transects placed next to the 24 herbivore exclusions installed at all main vegetation types of the END between 1992 and 2000 (17 of them in 1992-3), and monitored every spring and summer-autumn since their installation by the team led by R. Soriguer (who will collaborate actively in their analysis; see below for details). Each year, observers counted (and removed) the droppings present along 200x1 m transects, separately for each ungulate species, at the end of the growth season (September-October). 

Table 1: Location and characteristics of the herbivore-exclusion plots (and associated control plots) for which long-time series of plant standing crop are available to be used in the analyses of Tasks 1.2.2. and 2.1.1.

Position (GPS)
Dominant species
N36.99390 W6.44241
Herbaceous meadow (vera)
Choza Pelitos
N37.07739 W6.39863
N36.91508 W6.29282
Pozo Cornejo
N37.06459 W6.43063
Saltmarsh bulrush (S. maritimus)
Esquina Noguera
N37.04060 W6.42241
Saltmarsh bulrush (S. maritimus)
N36.98377 W6.39683
Somerset rush (Juncus subulatus)
Leo Biaggi - almajal
N36.97566 W6.32183
Saltworts + Somerset rush
Leo Biaggi - caño
N36.97586 W6.32304
Bulrush (S. litoralis)
Lucio del Rey - almajal
N36.91980 W6.34959
Lucio del Rey - lucio
N36.92227 W6.34745
Casa Matias
N37.01424 W6.32215
N37.01911 W6.36498
Bulrush (S. litoralis)
Cancela de Martinazo
N37.02831 W6.42930
Saltmarsh bulrush (S. maritimus)
Martinazo (viejo)
N37.03292 W6.43764
N37.00583 W6.50785
N37.03013 W6.43298
Pozo Almajal
N37.00765 W6.32695
N37.07502 W6.43317
Saltmarsh bulrush (S. maritimus) + spikerush (Eleocharis spp.)
Santa Olalla
N36.98216 W6.47814
Meadow (sandy shore of a large pond)
Caño Travieso
N37.00620 W6.31065
Somerset rush (Juncus subulatus) + saltworts
Veta del Palacio
N36.99211 W6.43368
Grassland (“veta”)
Veta Carrizosa
N36.97950 W6.40672
Grassland (“veta”)
Veta Marisma
N36.99299 W6.43445
Saltmarsh bulrush (S. maritimus)
N37.08515 W6.43321
Saltmarsh bulrush (S. maritimus) + spikerush (Eleocharis spp.)

1.2.3. To be able to refer these data to actual grazing pressure by the different herbivores, we will combine trimestral counts of droppings at the same transect with camera-trap photographs taken, at hourly intervals, at marked areas of vegetation placed next to such transects (see Table 1). Droppings will be collected to obtain deposition rates per interval and for fecal analysis (see Task 1.3.2.). Besides providing a calibration of the relation between dropping density and grazing pressure, data from camera traps will provide information on temporal (seasonal and diel) changes in such grazing pressure.

1.2.4. To obtain data on the spatial scale at which ungulates (from the five main species present at EBD: cattle, horse, red deer, fallow deer and wild boar) are exploiting the available plant resources, as well as the extent to which they are able to complement the availability of food items at different sites (different states within the EBD, and different areas within each state), we will analyze the movements of 10-20 individuals from each species using GPS trackers. Whenever possible, we will select individuals belonging to different social groups, so that their movements can be taken to represent that of the whole group (e.g. for cattle; Lazo 1995). Part of these data will be provided by R. Soriguer (EBD-CSIC) and J. Vicente (IREC-CSIC), who collected them in the course of previous projects focused on different aspect of the eco-epidemiology of tuberculosis (e.g. cattle and wild boar, in Barasona et al. 2014), thus expanding the time span and reducing the equipment costs of the current project. GPS collars will be installed within the first semester of the project, allowing for a tracking period of at least two years (though it will hopefully extent beyond such duration). Building on the availability of detailed (abiotic and biotic) environmental information from Doñana’s monitoring program (RBD-ICTS), data will be analyzed using the environmental-data automated track annotation (Env-DATA) system, which provides an easy-to-use platform for data acquisition, data transformation and integration, resampling, and interpolation (Dodge et al. 2013) that facilitates a comprehensive assessment of the basic question how movements are shaped by the environment (Nathan & Giuggioli 2013).

Task 1.3. Feeding requirements
- Achievement H3: Assessment of spatial and temporal changes in feeding requirements, associated to changes in food availability, food choice and digestion efficiency, completed (data collection and statistical tests).
- Deliverable E1: Scientific Article (together with Task 1.1. and 1.3.).

Previous studies have provided already global estimates of the feeding requirements of four ungulates addressed in this project (cattle, horse, red deer and fallow deer), based on available estimates of their energy and protein requirements (Soriguer et al. 2001). A similar procedure will be used to calculate the historical changes, as well as the spatial and temporal variation, in the feeding requirements of the populations of the five species of ungulates estimated in the previous Objective (Task 1.3.1.).

These studies acknowledge, however, that the estimates provided are subjected to substantial variation in diet across seasons (particularly, as the pasture dries in summer), among localities and across individuals. Such information is essential to evaluate the impact of competition among the different herbivore species, and their responses thereupon; to estimate the extent to which such species are able to respond to food shortages through changes in diet choice (e.g., complementing unpalatable or toxic food sources with palatable or undefended ones) and/or foraging areas (displacement); and to evaluate adequately the impact of the different herbivore species on the different  plant species (i.e., the actual drivers of changes in species composition under different grazing pressures, see below).

To obtain a more detailed description of herbivore diet composition choices and requirements for the different herbivore species, we will work at two different scales:

1.3.2. Fecal analysis: We will analyze the nutrient and energy content of the fecal samples collected in Task 1.2.3. and compare them with vegetation samples taken at the same moment, to estimate nutrient and energy uptake. These estimates will provide information on the intake rates required to meet energy and nutrient demands of the different ungulate species, and the different vegetation types and seasons.

1.3.3. Food analysis: We will register and analyze the diet of the individual herbivores tracked with GPS collars (Task 1.2.4) by means of video-recordings obtained by animal-borne video-cameras (e.g. Loyd et al. 2013). These systems have a proven accuracy to provide information on numerous aspects of the behavior of elusive species, including the selection of food item to the level of individual plant species (e.g. Thompson et al. 2012). In largely mobile and elusive animal, the use of such systems has been limited by its high cost. In our case, however, the relatively easy access to the domestic herbivores will allow us to use relative inexpensive systems that must be renewed over shorter periods of time (e.g. 7-10 days for the Kitty Cams Project, For more elusive animals (deer and wild boar), we will resort to more expensive equipment with prolonged battery life and detachable collars (to recover the images without having to recapture the individual). Cameras will be deployed, for periods of 1-2 weeks, at four different moments in the year (autumn, winter, spring and summer).

Ob.2. Estimate the effects of rainfall level and herbivore pressure on the productivity and standing crop of the vegetation of Doñana

Task 2.1. Analyze historical data on the productivity and standing crop of the vegetation of Doñana.
- Achievement H4: Assessment of historical changes in the productivity and standing crop of the vegetation of Doñana, and the effect of rainfall and grazing pressure thereupon, completed (data collection and statistical tests).
- Achievement H5: Collection of a dataset for ground-truthing the standing-crop estimates obtained from RS data (Ob.3) completed.
- Deliverable E2: Scientific Article (together with Task 2.2.).

In the course of previous studies and management interventions, a considerable number of exclusion plots (and associated control areas) have been installed and monitored at the different vegetation/landscape units of the END. We will analyze the available data of those which have been monitored in previous years, and monitor those that will be made available during the course of this year, as follows:

2.1.1. Analysis of historical data on community composition and plant standing crop from 24 (excluded and control) plots installed at all main vegetation types of the END between 1992 and 2000, and monitored every spring and summer-autumn since their installation by the team led by R. Soriguer (who will collaborate actively in their analysis (see Table 1). Monitoring included record of the cover (per species) and biomass yield (all species pooled) inside and outside the exclusion plots. Cover and biomass yield (aboveground biomass, clipped at ground level) were estimated at the end of each growth season (September-October) in 1x1 m squares (N=8 within the exclusion and N=20 along 200-m transects placed outside the exclusion).

2.1.2. Monitoring the community composition and plant standing crop of 9 plots that will be installed at the end of 2016, in the course of Life+ Project ADAPTAMED. These plots will aim at facilitating the recruitment of large shrubs and trees, and such recruitment will be monitored twice a year by the ADAPTAMED team. We will be allowed to join in and monitor the two variables mentioned above, as well as the analyses of grazing pressure and dropping production described in Objectives 1.2.3. and 1.3.2. The use of this plots will fill the main gap of those for which historical data are available (task 2.1.1.), since they are located at shrubland areas within three different livestock management areas (thus largely different grazing pressure) of the END. Cover and biomass yield will be estimated as in task 2.1.1.

2.1.3. Monitoring the community composition and peak standing crop at 10 additional sites per vegetation unit (as defined in Obj.3) during the first two years of the project, with the aim of providing data to validate the remote sensing models developed in Obj.3. Cover and biomass yield will be estimated as in task 2.1.1.

Task 2.2. Experimental determination of the effect of precipitation and herbivory on plant productivity and standing crop
- Achievement H6: Assessment of the effect of precipitation and herbivory on plant productivity and standing crop  (experiment and statistical tests).
- Deliverable E2: Scientific Article (together with task 2.1.)

To obtain experimental estimates of the separate and combined effect of precipitation and herbivory on the productivity of the Doñana vegetation. For this purpose, we will use two vegetation types, chosen to represent situations with low and high plant diversity: the “marisma de castañuela”, dominated by a single species, Scirpus maritimus; and the highly-diverse pastures of the ecotone between the marsh and the sand, the “vera”. In both cases, we will cross factorially two types of treatments with four levels each: (i) Rainfall: four levels corresponding to the mean, median, upper (75%) and lower (25%) quartile of the historical rainfall record. The distribution of rainfall over time will also reproduce the average distribution observed in the historical record (for the subset of year in which a precipitation equal, with 10%, to the chosen level of annual rainfall). (ii) Herbivory: Removal of the upper 0%, 33%, 66%and 100% part of the plant (to the nearest cm of the plant’s length) present at each monthly interval. The proportion of biomass removed will be weighed to estimate total plant production at the end of the experiment. A subsample of the plants from each treatment will be harvested each month, to estimate biomass yield, biomass allocation (to leaves, shoots, rhizomes, roots and, if present, tubers). The experiment will take place at the experimental facilities (mesocosm facility) and field plots (provided with herbivore-exclusion fences and water supply) of the Doñana Biological Reserve (RBD-ICTS).

Ob.3. Reconstruct the long-term changes in productivity and standing crop of the Doñana vegetation, using remote-sensing tools
- Achievement H7: Development, calibration and validation of RS models estimating the productivity and standing crop of the vegetation of Doñana.
- Deliverable E3: Scientific Article.

We will combine remote sensing data, consisting of NDVI estimated derived from MODIS satellite images from the study area, with biomass production data obtained in the previous objective (Ob.2), to produce models allowing for the estimation of plant biomass (standing crop) and productivity (changes in plant standing crop) from such images. These model will be used to (i) model the seasonal cycle of biomass production, and its variation across years and its relation to total biomass yield, (ii) model the effect of key environmental factors (such as rainfall, topography and grazing pressure) on such cycles, and (iii) reconstruct historical changes in biomass yield (last three decades) and evaluate its potential drivers (e.g. changes in grazing pressure). For this purpose, we will proceed in
three steps:
Task 3.1. Obtain long-term series of NDVI for the different vegetation types of the END and fit them to existing models based on key environmental factors (such as rainfall, topography and grazing pressure).

This task will be based on the vegetation units defined and mapped at the “Cartografía de los Sistemas de Vegetación del Parque Nacional de Doñana 1:10000” (2014). This is the last vegetation map published. It is based in the interpretation of orthophotos from 2009 and it incorporates previously published vegetation maps.

For each of the main vegetation units for which biomass yield data are available (see Table 1), we will produce a NDVI temporal curve at 10-16 groups of 3x3 pixels (‘sites’ hereafter). These sites will include those for which biomass data are available (N=1-6), which will be used for cross-calibration; plus 10 additional sites, distributed as to maximize variation in current plant productivity (estimated from a classification of average NDVI values during the last five years), which will be used for validation (see below).

Data will be derived from MODIS satellite images (spatial resolution of 250 m, temporal resolution: 7 days) from 2000 until the last image available at the time of task execution. The series that we will use is already processed in form of NDVI values provided by BOKU Vienna (Vuolo, Mattiuzzi, Klisch, & Atzberger, 2012).
NDVI temporal curves characterize the development and photosynthetic activity of the green vegetation. Firstly, we will derive phenological parameters of the curves in two steps. We will use Timesat, a software package specifically developed to analyze time-series of satellite data (Jonsson & Eklundh, 2004), to analyze the time series derived from MODIS images (since Timesat does not accept irregular temporal series). In a second step, we will compare the interannual variation in phenological parameters across vegetation units, to evaluate their reliability as food resource for herbivores; and will evaluate the effect of different environmental variables (such as precipitation, inundation probability and/or grazing pressure) on such variation.

Task 3.2. Calibrate and validate the predictions of the model, using the available data of plant biomass obtained in Task 1.2.

Phenological curves fitted to NDVI values will be used to obtain different estimates of plant standing crop (based, e.g., on the peak values or the area under the curve). Separate subsets of these values will be used for calibration and validation, as follows:

3.1.1.        At the sites where biomass-yield data are available (N=1-3), annual estimates derived from NDVI curves (N=20-30 per site) will be compared to ground-truth values and their relationship modelled using Generalized Linear (Mixed) Models. Departing from full models with random intercept and slopes, we will select the most parsimonious model based on the AIC scores.

3.1.2.        Based on the previous calibration, we will calculate the biomass yield of the remaining sites (N=17-19) for the first two years of the project, and compare these values to field observations of peak biomass yield taken at these two years.

Task 3.3. Extrapolate the available estimates to the rest of the END.

Based on the parameter values obtained in the previous sections, and the NDVI values derived from MODIS images, we will produce maps of plant standing crop across the whole END. These maps will be used to (i) estimate the carrying capacity of the plant’s vegetation for herbivores, and its recent changes (last 2-3 decades), and (ii) estimate the interaction between different descriptors of space use and grazing pressure by ungulates, and the spatial variation in the availability of food resources (plant standing crop).

Ob.4. Develop models of the relationship between rainfall, grazing pressure and plant standing crop at the END, and use them to evaluate different management scenarios under current and forecasted climate conditions.

Task 4.1. Develop models of the effect of rainfall and grazing on plant standing crop
- Achievement H8: Development, calibration and validation of spatially-explicit models of the relationship between rainfall, grazing pressure and plant standing crop at the END.
- Achievement H9: Collaborative design, generation and evaluation of scenarios reflecting different management strategies and forecasted changes in climatic conditions.
- Deliverable E4: Scientific Article.
- Deliverable E5: Adaptive management strategy. (The achievement of this deliverable is contingent on achieving the necessary levels of trust and cooperation among managements and stakeholders.)

Based on the results of previous objectives, we will develop spatially-explicit, process-based models of the relationship between rainfall, grazing pressure and plant standing crop. We will model the behavior of the different vegetation units of the END separately, while allowing for the ungulates to move freely among these units (within the restrictions imposed by the physical barriers associated to the different management units, see above). The model will incorporate simple demographic models for wild ungulates, while the abundances of domestic herbivores will be considered as fixed (i.e. as determined by the management regime).

The main state variable of the model will be plant standing crop. This is the main descriptor of plant production addressed throughout the project, owing largely to its availability from historical observations. However, this variable is actually the result of the dynamic interplay between plant growth and plant consumption by herbivores. To be able to derive estimates of plant biomass available for herbivore consumptions, the model will simulate these two processes separately. To evaluate the reliability of these predictions, we will calibrate and validate the model using the estimates of plant standing crop obtained in Ob.3 (Task 3.3.), both separately for each vegetation unit and pooled across them. Separated subsets for calibration and validation will be obtained in two different ways: (i) by splitting the complete time series in two (10-15 years each), and (ii) by randomly sampling separate sets of observation units (3x3 pixels) for calibration and validation, within each year of the time series (following autocorrelation analysis to ensure independence of the two datasets).

Task 4.2. Generate and evaluate different management scenarios under current and forecasted climate conditions (climate change)

The model developed in 5.2. will be used to evaluate different scenarios arising from the factorial combination of different management strategies and different climate conditions (both current and forecasted). The development and evaluation of the different management scenarios will be undertaken in close collaboration with the END managers and key stakeholders (e.g. local rancher associations, environmental NGOs), using participatory techniques in which the research team has already ample experience (e.g. Méndez et al. 2012, Santamaría et al. 2013). These interactions will be structured in three different workshops, respectively addressing (i) the purpose and structure of the model, (ii) model development and preliminary model simulations, (iii) final, interactive simulations. If possible (i.e., if the level of trust and collaboration raised during the participatory modelling phase allows for it), we will aim at the elaboration of a (draft) adaptive management plan for the domestic and wild ungulates of the END to be a result of the third workshop.

Work plan

Workteam members
Year 1
Year 2
Year 3
Task 1.1.Historical data on herbivore abundance
Task 1.2. Spatial distribution and individual movements

1.2.1. Analyze monitoring data
1.2.2. Analyze dropping counts
1.2.3. Camera traps
1.2.4. GPS collars
Task 1.3. Feeding requirements

1.3.1. Literature analysis
1.3.2. Fecal analysis
1.3.3. Food analysis
Task 2.1. Historical data on the plant standing crop

2.1.1. Analyze historical data
2.1.2. Survey shrubland plots
2.1.3. Survey additional sites
Task 2.2. Experiment on effects of precipitation and herbivory

Task 3.1. NDVI models
Task 3.2. Calibrate and validate model predictions
Task 3.3. Maps of plant standing crop
Task 4.1. Develop dynamic models of plant standing crop
Task 4.2. Generate and evaluate different management scenarios



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The main contribution expected from this project will be the development of a knowledge base, long-term criteria and modelling tools for the management of free-ranging livestock in Mediterranean protected areas with high climatic variability, taking the Doñana Protected Area (Espacio Natural Doñana, END) as a case study. The project impact should therefore span from general science, where it will provide insight into cutting-edge questions such as the spatial factors underpinning resource use by herbivores and the role of landscape heterogeneity as a buffer against the crossing of tipping points leading to environmental degradation; to applied research, where it will provide specific tools and guidelines for the management of free-ranging livestock-production system, which should be readily applicable to Doñana and, should they prove useful, could be extrapolated to other areas with similar characteristics. More importantly, it will provide elements for the proactive management of such exploitations in the face of climate change, increasing the resilience of socio-natural system against expected changes associated to it. The participatory approach proposed during the modeling phase will increase the likelihood of a speedy transfer of he project results, and will hopefully introduce elements of social innovation in a management system that has traditionally relied quite exclusively on specialist from the fields of natural sciences append engineering.
Besides several deliverables directly applicable to knowledge transfer into policy and management (RS and dynamic models, collaborative adaptive management strategy), the project will also aim at achieving a high-quality scientific output, in the form of (i) four publications in high impact journals (such as Ecography, Ecology, J Ecol, J Appl Ecol, Conserv Biol or Ecol Modelling, where we have published extensively in the past) - two of them, if funding allows, in open-access journals to increase their visibility; (ii) one article summarizing the results in a science journal for the general public; (iii) one national (Spanish Association of Terrestrial Ecology AEET) and one international conference (Ecological Society of America, British Ecological Society or INTECOL); and (iv) two presentations for practitioners (Park Managers), stakeholders (livestock-breeder associations, environmental NGOs) and the general public (e.g. Science Week, technical and secondary schools).
Outreach towards academics, practitioners and more general public will be also achieved through the development of a project web page, hosted at the general page of the EBD-CSIC, where all project results (including modeling products) will be made publicly available. Entries to this page will be disseminated using other social media, through the accounts managed by the outreach team of EBD-CSIC.


The IP and other researchers of the work team has formed a high number of researchers that have obtained their PhD thesis, MSc degrees or senior graduate projects. In particular, Luis Santamaría has supervised five MSc Theses and seven PhD Thesis. The later include those defended by Jörn Pilon at Wageningen University (The Netherlands) in 2002; Helen Hangelbroek, at Nijmegen University (The Netherlands) in 2004; Ainhoa Magrach, at the University of Santiago de Compostela (Spain), in 2011; Iris Charalambidou, at Utrecht University (The Netherlands), in 2012; Lucía Latorre, at the University of Santiago de Compostela (Spain), in 2013; Duarte Viana at the Pablo de Olavide University ( Spain) and Pablo Fernández Méndez, at the University of the Balearic islands (Spain). All these PhD theses obtained outstanding marks and resulted in high quality scientific papers.

The interdisciplinarity and the high quality of the working team are a guarantee of a good environment for the formation of new researchers. At the same time, the IP and sole member of the research team (Luis Santamaría) will be able to provide fully-committed supervision to the PhD student requested, since his only commitment at present is the co-supervision of one PhD researcher, who will be in the third year of her work by 2017. The project results will represent a suitable departure point from which the PhD student will be able to add an in-depth assessment of the effects of ungulates on other elements of the Doñana ecosystem, having from the start the freedom to select a topic of his/her own and the opportunity to benefit from the scientific quality, interdisciplinarity and internationality of the work team in developing such topic.