An introduction to Mediterranean plant ecology. The course covers the main topics on basic Ecology (abiotic factors, interactions, disturbance, succession, diversity) while providing a comprehensive picture of the current main ecological problems, such as invasions, global warming, loss of biodiversity, habitat restoration, etc., all in the context of Mediterranean ecology. Three field trips complete the course, with field work directly related to the topics offered in the classroom, including field sampling, collection of data, and a basic in situ analysis, in order to draw conclusions.
Prerequisites:
One semester of college-level biology or consent of instructor.
Method of presentation:
Lectures, case studies, group discussions with student presentations, and three field study visits. For each unit, we will provide a general outline of the subject, and later the topic will be approached and discussed with specific texts that the students must read (readings indicated for each unit, see below). List of readings may be slightly modified to incorporate new scholarship on the subject.
Field study:
Field trip 1. North of Granada province. Badland area of Guadix-Baza basin. Field work will be related to plant- plant interaction and facilitation. We will analyze the spatial relationship between two common species in the area, a shrubs (Retama sphaerocarpa) and a scrub (Artemisia campestris). After data collection, the students will calculate a few parameters with the help of a pocket calculator and will test whether these two species are positively or negatively associated. The results will be discussed in the context of the competition-facilitation gradient. Additional aspects of dry ecosystems will be discussed on this field trip.
Readings: Pugnaire F.I., Haase P. and Puigdefábregas J. (1996). 1420-1426.
Rodríguez-Echevarría S. and Pérez-Fernández M.A. (2003). 807-814.
Field trip 2. Sierra de Huétor Natural Park area, field work (morning) and visit to the park (evening). The field work will be related to disturbance and succession (fire ecology), including aspects of plant dispersal and plant defense against herbivores. The dynamics of regeneration after fire will be analyzed in a burnt area, considering the two main disturbance agents: fire (high intensity, low frequency) and herbivores (low intensity, high frequency). Students will sample the recruitment of the main shrubs and trees of the forests, either sprouters or seeders, and will compare data with other sampling in adjacent, non-burned area. With the data collected and a few simple calculations the successional trends of the community will be discussed, and their consequences for management. During the evening, the area will be visited, with special attention to the effect of herbivory on plant traits (e.g. spinescence), and dispersal of juniper “fruits” as well as acorns by birds.
Readings: Ojeda F., Marañón T. and Arroyo J. (1996). 191-198.
Pausas J.G., Ribeiro E. and Vallejo R. (2004). 251-259.
Field trip 3. La Cortijuela Botanical Garden area, Sierra Nevada National Park. A field trip where a great amount of research is currently being performed in relation to Mediterranean plant ecology. Aspects of seed dispersal, herbivory, seedling recruitment and establishment, biotic and abiotic factors determining forest regeneration, etc., will be discussed while visiting the experimental areas.
Readings: Castro J., Zamora R., Hódar J.A. and Gómez J.M. (2004). 266-277.
Gómez-Aparicio L., Zamora R. and Gómez J.M. (2005). 195-206.
Required work and form of assessment:
Oral and/or written reports on weekly assigned readings 20%
Reports on each field study visit 35% total
Mid-term examination 20%
Final examination 25%
Attendance to all lectures and field trips is mandatory.
content:
Unit 1. Setting the scene: Determinants of Mediterranean-type ecosystems
The physical scenario: recent geological history of the Mediterranean Region. Topography. Distribution of main mountain ranges and effect on local climatic conditions.
Abiotic factors: Climate, light, and soils. Determinants of climate in the Mediterranean region. Vicissitudes during the Pleistocene. Current climatic conditions. Annual regime of precipitation and temperature: seasonal uncoupling of conditions for growth. Light and soil moisture as ecological factors: intensity and spatial pattern of distribution.
Nutrients. Source of nutrients. Spatial and temporal pattern of variation. Decomposition and mineralization of nutrients and plant growth and recruitment. Plant-soil-microbial interactions. Association with other organisms (mycorrhizas).
Case studies:
How to cope with arid environments.
The uncoupling of soil moisture, light and temperature determining the establishment of a tree species.
Readings:
Blondel J. and Aronson J. (1999). Chapter 1.
Puigdefábregas J. and Pugnaire F.I. (1999). in Pugnaire F.I. and Valladares F. (eds.), Handbook of functional plant ecology. Pp. 381-405.
Castro J., Zamora R., Hódar J.A. and Gómez J.M. (2005).
Unit 2. Life-history traits and functional attributes in Mediterranean-type ecosystems
Evergreenness and sclerophylly. Life forms and their relation to the habitat. Flowering patterns and pollination systems: anemogamous versus zoogamous strategies. Seed dormancy as a way of resistance. Adaptations to water scarcity. Adaptations to nutrient deficits. Adaptations to high light intensity and ultraviolet radiation. Adaptations to high herbivore pressure. Plant acclimation to environmental conditions.
Case studies:
Herbivory induces spinescence in tree leaves.
Life-history traits such as root:shoot ratios and seed size determine seedling survival and may affect community composition.
Readings:
Joffre R., Rambal S. and Damesis C. (1999). in Pugnaire F.I. and Valladares F. (eds.), Handbook of functional plant ecology. Pp. 347-380.
Lloret F., Casanovas C. and Peñuelas J. (1999). 210-216.
Obeso J.R. (1997). 149-156.
Unit 3. Interactions among plants: competition, facilitation and parasitism
Plant competition in Mediterranean-type ecosystems: Magnitude and relevance. Allelopathy. From competition to facilitation: two extremes of a gradient. Mechanisms of facilitation. Plants that parasite plants.
Case studies:
Taking advantage of facilitation for forest reforestation.
The impact of parasitic plants also depends on other factors: the effect may be positive under high herbivore pressure.
Readings:
Callaway R.M. and Pugnaire F.I. (1999). in Pugnaire F.I. and Valladares F. (eds.), Pp. 623-648 rk.
Gómez J.M. (1994). 97-106.
Gómez-Aparicio L., Zamora R., Gómez J.M., Hódar J.A., Castro J. and Baraza E. (2004). 1128-1138.
Unit 4. Interactions with animals
Animals as predators: Vertebrate herbivores, invertebrate herbivores, and seed predators. Defense mechanisms in plants: Chemical defense, physical defense, and associational resistance. Animals as pollinators and animals as seed dispersers: From predation to mutualism.
Case studies:
Combining facilitation with animal dispersal: Two critical steps that act synergetically in yew regeneration.
Jays and dispersal of acorns: Certain predation may result positive if the plant finally derives some benefit.
Associational resistance versus shared dooms: The outcome of the interaction is also context-dependent.
Readings:
Baraza E., Zamora R. and Hódar J.A. (2005).
García D., Zamora R., Hódar J.A, Gómez J.M. and Castro J. (2001). 31-38.
Gómez J.M. (2003). 573-584.
Zamora R., Hódar J.A. and Gómez J.M. (1999). In Pugnaire F.I. and Valladares F. (eds.), Handbook of functional plant ecology. Pp. 677-718.
Unit 5. Disturbance and succession in Mediterranean plant communities
Disturbance regimes: magnitude, frequency, and recurrence. Herbivores as a source of disturbance in
Mediterranean-type ecosystems. Fire as a major disturbance in Mediterranean type-ecosystems. Fire ecology in Mediterranean-type ecosystems. Coping with fire: resprouters and seeders.
Succession. Mechanisms of succession. Synchronic and diachronic models. Fire, man, and their historical impact on succession dynamics. The concept of climax and critics received. Major plant communities of Andalusia.
Case studies:
Man, fire, and community composition in the Mediterranean area: consequences of millennia of human presence.
Fire and herbivores as determinants of succession and mature community composition. Climatic conditions are not the only important factor.
Readings:
Bond W.J. and Keeley J.E. (2005). 387-394.
Carrión J.S., Yll E., Willis K.J. and Sánchez P. (2004). 219-236. Whelan R.J. (1995). Chapter 2.
Unit 6. Biodiversity of the Mediterranean region
Measurements of biodiversity. Hot spots of biodiversity. Present day biodiversity in the Mediterranean region. Rate of endemisms. Paleohistorical causes of high biodiversity in the Mediterranean region. Patterns of diversity: effect of altitude, latitude, and habitat productivity. Biodiversity, productivity, and stability. Redundancy and biodiversity. The decline of biodiversity in recent times. Pros and cons of human impact.
Case studies:
Set of interactions that affect biodiversity: from positive to negative effects of grazing.
Change in climatic conditions may impact diversity in Mediterranean-type ecosystems.
Readings:
Lloret F., Peñuelas J. and Estiarte M. (2004). 248-258.
Myers N., Mittermeler R.A., Mittermeler C.G., da Fonseca G.A.B. and Kent J. (2000). 853-858. Osem Y., Perevolotsky A. and Kigel J. (2004). 936-946.
Whitton J. & Rajakaruna N. (2001). in Levin S.A. (ed.) Encyclopedia of biodiversity, Vol. 4. Pp. 621-630.
Unit 7. Global change and Mediterranean-type ecosystems
Causes of global change. Consequences of global change on climatic parameters: predictions for the Mediterranean area. Consequences of global change on the biota: mechanism of action. Biota with higher risk under global change.
Biological invasions. Consequences of biological invasions for local biota. Habitat invasiveness. Traits related to species invasiveness. Overall consequences on biodiversity.
Case studies
Global warming and loss of synchrony in ecosystems: a caterpillar threatens Scots pine forests in the Mediterranean mountain.
Plant stoichiometry, herbivores diet, and human nutrition: an example of how global change affects human beings.
General patterns of invasiveness: trying to predict models for ecosystem management and restoration.
Readings:
Dunne J.A. and Harte J. (2001). in Levin S.A., Encyclopedia of Biodiversity, vol. 3. Pp. 277-293
Loladze I. (2002). 457-461.
Hódar J.A, Castro J. and Zamora R. (2003). 123-129.
Lloret F., Medail F., Brundu G., Camarda I., Moragues E., Rita j., Lambdon P. and Hulme P.E. (2005). 512-520.
Perrings C., Dehnen-Schmutz K., Touza J. and Williamson M. (2005). 212-215.
Required readings:
Baraza E., Zamora R. and Hódar J.A. (2005). Conditional outcomes in plant-herbivore interactions: neighbors matters. Oikos (in press).
Bond W.J. and Keeley J.E. (2005). Fire as a global “herbivore”: the ecology and evolution of flammable ecosystems. Trends in Ecology and Evolution 7: 387-394.
Blondel J. and Aronson J. (1999). Biology and wildlife of the Mediterranean region. Oxford University Press, U.K. Chapter 1.
Callaway R.M. and Pugnaire F.I. (1999). Facilitation in plant communities. Pp. 623-648 in: Pugnaire F.I. and Valladares F. (eds.), Handbook of functional plant ecology. Marcel Dekker, New York.
Carrión J.S., Yll E., Willis K.J. and Sánchez P. (2004). Holocene forest history of the eastern plateaux in the Segura Mountains (Murcia, southeastern Spain). Review of Paleobotany and Palynology 132: 219-236.
Castro J., Zamora R., Hódar J.A. and Gómez J.M. (2004). Seedling establishment of a boreal tree species (Pinus sylvestris) at its southernmost distribution limit: consequences of being in a marginal Mediterranean habitat. Journal of Ecology 92: 266-277.
-----. (2005). Alleviation of summer drought boost establishment success of Pinus sylvestris in a Mediterranean mountain: an experimental approach. Plant Ecology (in press).
Dunne J.A. and Harte J. (2001). Greenhouse effect. Pp. 277-293 en: Levin S.A., Encyclopedia of Biodiversity, vol. 3. Academic Press.
García D., Zamora R., Hódar J.A, Gómez J.M. and Castro J. (2001). Yew (Taxus baccata L.) regeneration is facilitated by fleshy-fruited shrubs in Mediterranean environments. Biological Conservation 95: 31-38.
Gómez J.M. (2003). Spatial patterns in long-distance dispersal of acorns by jays in a heterogeneous landscape. Ecography 26: 573-584.
-----.(1994). Importance of direct and indirect effects in the interaction between a parasitic plant (Cuscuta epithymum) and its host plant (Hormathophylla spinosa). Oikos 71: 97-106.
Gómez-Aparicio L., Zamora R., Gómez J.M., Hódar J.A., Castro J. and Baraza E. (2004). Applying plant positive interactions to reforestation in Mediterranean mountains: A meta-analysis of the use of shrubs as nurse plants. Ecological Applications 14: 1128-1138.
Gómez-Aparicio L., Zamora R. and Gómez J.M. (2005). The regeneration status of the endangered Acer opalus subsp granatense throughout its geographical distribution in the Iberian Peninsula. Biological Conservation 121: 195-206.
Hódar J.A, Castro J. and Zamora R. (2003). Pine processionary caterpillar Thaumetopoea pityocampa as a new threat for relict Mediterranean Scots pine forests under climatic warming. Biological Conservation 110: 123- 129.
Joffre R., Rambal S. and Damesis C. (1999). Functional attributes in Mediterranean-type ecosystems. Pp. 347-380 in: Pugnaire F.I. and Valladares F. (eds.), Handbook of functional plant ecology. Marcel Dekker, New York.
Lloret F., Medail F., Brundu G., Camarda I., Moragues E., Rita j., Lambdon P. and Hulme P.E. (2005). Species attributes and invasion success by alien plants on Mediterranean islands. Journal of Ecology 93: 512-520.
Lloret F., Peñuelas J. and Estiarte M. (2004). Experimental evidence of reduced diversity of seedlings due to climate modification in a Mediterranean-type community. Global Change Biology 10: 248-258.
Lloret F., Casanovas C. and Peñuelas J. (1999). Seedling survival of Mediterranean shrubland species in relation to root:shoot ratio, seed size and water and nitrogen use. Functional Ecology 13: 210-216.
Loladze I. (2002). Rising atmospheric CO2 and human nutrition: toward globally imbalanced plant stoichiometry? Trends in Ecology and Evolution 17: 457-461.
Myers N., Mittermeler R.A., Mittermeler C.G., da Fonseca G.A.B. and Kent J. (2000). Biodiversity hotspots for conservation priorities. Nature 403: 853-858.
Obeso J.R. (1997). The induction of spinescence in European holly leaves by browsing ungulates. Plant Ecology 129: 149-156.
Osem Y., Perevolotsky A. and Kigel J. (2004). Grazing effect on diversity of annual plant communities in a semi-arid rangeland: interactions with small-scale spatial and temporal variation in primary productivity. Journal of Ecology 90: 936-946.
Ojeda F., Marañón T. and Arroyo J. (1996). Postfire regeneration of a Mediterranean heatland in southern Spain. International Journal of Wildland Fire 6: 191-198.
Pausas J.G., Ribeiro E. and Vallejo R. (2004). Post-fire regeneration variability of Pinus halepensis in the eastern Iberian Peninsula. Forest Ecology and Management 203: 251-259.
Perrings C., Dehnen-Schmutz K., Touza J. and Williamson M. (2005). How to manage biological invasions under globalization. Trends in Ecology and Evolution 20: 212-215.
Pugnaire F.I., Haase P. and Puigdefábregas J. (1996). Facilitation between higher plant species in a semiarid environment. Ecology 77: 1420-1426.
Puigdefábregas J. and Pugnaire F.I. (1999). Plant survival in arid environments. Pp. 381-405 in: Pugnaire F.I. and Valladares F. (eds.), Handbook of functional plant ecology. Marcel Dekker, New York.
Rodríguez-Echevarría S. and Pérez-Fernández M.A. (2003). Soil fertility and herb facilitation mediated by Retama sphaerocarpa). Journal of Vegetation Science 14: 807-814.
Whelan R.J. (1995). The ecology of fire. Cambridge University Press, Cambridge, U.K. Chapter 2.
Whitton J. & Rajakaruna N. (2001). Plant biodiversity, overview. Pp. 621-630 in: Levin S.A. (ed.) Encyclopedia of biodiversity, Vol. 4. Academic Press, San Diego.
Zamora R., Hódar J.A. and Gómez J.M. (1999). Plant-herbivore interaction: beyond a binary vision. Pp. 677- 718 in: Pugnaire F.I. and Valladares F. (eds.), Handbook of functional plant ecology. Marcel Dekker, New York.
Brief Biography of Instructor:
Dr. Jorge Castro is a Professor of Ecology at the Universidad de Granada, and a member of the UGR's Terrestrial Ecology Research Group, which states as its goals, 1) to broaden knowledge of key ecological processes in Mediterranean ecosystems, 2) to develop conceptual tools and methodologies to improve the conservation and management of natural resources, and 3) to promote the transfer of research results to the management of protected areas, disseminating this information through training and advisory activities. Dr. Castro has a very extensive record of publications and presentations in the field. More information available at www.ugr.es/~rnm220/.
Mediterranean Ecosystems
An introduction to Mediterranean plant ecology. The course covers the main topics on basic Ecology (abiotic factors, interactions, disturbance, succession, diversity) while providing a comprehensive picture of the current main ecological problems, such as invasions, global warming, loss of biodiversity, habitat restoration, etc., all in the context of Mediterranean ecology. Three field trips complete the course, with field work directly related to the topics offered in the classroom, including field sampling, collection of data, and a basic in situ analysis, in order to draw conclusions.
One semester of college-level biology or consent of instructor.
Lectures, case studies, group discussions with student presentations, and three field study visits. For each unit, we will provide a general outline of the subject, and later the topic will be approached and discussed with specific texts that the students must read (readings indicated for each unit, see below). List of readings may be slightly modified to incorporate new scholarship on the subject.
Field trip 1. North of Granada province. Badland area of Guadix-Baza basin. Field work will be related to plant- plant interaction and facilitation. We will analyze the spatial relationship between two common species in the area, a shrubs (Retama sphaerocarpa) and a scrub (Artemisia campestris). After data collection, the students will calculate a few parameters with the help of a pocket calculator and will test whether these two species are positively or negatively associated. The results will be discussed in the context of the competition-facilitation gradient. Additional aspects of dry ecosystems will be discussed on this field trip.
Readings:
Pugnaire F.I., Haase P. and Puigdefábregas J. (1996). 1420-1426.
Rodríguez-Echevarría S. and Pérez-Fernández M.A. (2003). 807-814.
Field trip 2. Sierra de Huétor Natural Park area, field work (morning) and visit to the park (evening). The field work will be related to disturbance and succession (fire ecology), including aspects of plant dispersal and plant defense against herbivores. The dynamics of regeneration after fire will be analyzed in a burnt area, considering the two main disturbance agents: fire (high intensity, low frequency) and herbivores (low intensity, high frequency). Students will sample the recruitment of the main shrubs and trees of the forests, either sprouters or seeders, and will compare data with other sampling in adjacent, non-burned area. With the data collected and a few simple calculations the successional trends of the community will be discussed, and their consequences for management. During the evening, the area will be visited, with special attention to the effect of herbivory on plant traits (e.g. spinescence), and dispersal of juniper “fruits” as well as acorns by birds.
Readings:
Ojeda F., Marañón T. and Arroyo J. (1996). 191-198.
Pausas J.G., Ribeiro E. and Vallejo R. (2004). 251-259.
Field trip 3. La Cortijuela Botanical Garden area, Sierra Nevada National Park. A field trip where a great amount of research is currently being performed in relation to Mediterranean plant ecology. Aspects of seed dispersal, herbivory, seedling recruitment and establishment, biotic and abiotic factors determining forest regeneration, etc., will be discussed while visiting the experimental areas.
Readings:
Castro J., Zamora R., Hódar J.A. and Gómez J.M. (2004). 266-277.
Gómez-Aparicio L., Zamora R. and Gómez J.M. (2005). 195-206.
Attendance to all lectures and field trips is mandatory.
Unit 1. Setting the scene: Determinants of Mediterranean-type ecosystems
The physical scenario: recent geological history of the Mediterranean Region. Topography. Distribution of main mountain ranges and effect on local climatic conditions.
Abiotic factors: Climate, light, and soils. Determinants of climate in the Mediterranean region. Vicissitudes during the Pleistocene. Current climatic conditions. Annual regime of precipitation and temperature: seasonal uncoupling of conditions for growth. Light and soil moisture as ecological factors: intensity and spatial pattern of distribution.
Nutrients. Source of nutrients. Spatial and temporal pattern of variation. Decomposition and mineralization of nutrients and plant growth and recruitment. Plant-soil-microbial interactions. Association with other organisms (mycorrhizas).
Case studies:
Readings:
Unit 2. Life-history traits and functional attributes in Mediterranean-type ecosystems
Evergreenness and sclerophylly. Life forms and their relation to the habitat. Flowering patterns and pollination systems: anemogamous versus zoogamous strategies. Seed dormancy as a way of resistance. Adaptations to water scarcity. Adaptations to nutrient deficits. Adaptations to high light intensity and ultraviolet radiation. Adaptations to high herbivore pressure. Plant acclimation to environmental conditions.
Case studies:
Readings:
Unit 3. Interactions among plants: competition, facilitation and parasitism
Plant competition in Mediterranean-type ecosystems: Magnitude and relevance. Allelopathy. From competition to facilitation: two extremes of a gradient. Mechanisms of facilitation. Plants that parasite plants.
Case studies:
Readings:
Unit 4. Interactions with animals
Animals as predators: Vertebrate herbivores, invertebrate herbivores, and seed predators. Defense mechanisms in plants: Chemical defense, physical defense, and associational resistance. Animals as pollinators and animals as seed dispersers: From predation to mutualism.
Case studies:
Readings:
Unit 5. Disturbance and succession in Mediterranean plant communities
Disturbance regimes: magnitude, frequency, and recurrence. Herbivores as a source of disturbance in
Mediterranean-type ecosystems. Fire as a major disturbance in Mediterranean type-ecosystems. Fire ecology in Mediterranean-type ecosystems. Coping with fire: resprouters and seeders.
Succession. Mechanisms of succession. Synchronic and diachronic models. Fire, man, and their historical impact on succession dynamics. The concept of climax and critics received. Major plant communities of Andalusia.
Case studies:
Readings:
Unit 6. Biodiversity of the Mediterranean region
Measurements of biodiversity. Hot spots of biodiversity. Present day biodiversity in the Mediterranean region. Rate of endemisms. Paleohistorical causes of high biodiversity in the Mediterranean region. Patterns of diversity: effect of altitude, latitude, and habitat productivity. Biodiversity, productivity, and stability. Redundancy and biodiversity. The decline of biodiversity in recent times. Pros and cons of human impact.
Case studies:
Readings:
Unit 7. Global change and Mediterranean-type ecosystems
Causes of global change. Consequences of global change on climatic parameters: predictions for the Mediterranean area. Consequences of global change on the biota: mechanism of action. Biota with higher risk under global change.
Biological invasions. Consequences of biological invasions for local biota. Habitat invasiveness. Traits related to species invasiveness. Overall consequences on biodiversity.
Case studies
Readings:
Dr. Jorge Castro is a Professor of Ecology at the Universidad de Granada, and a member of the UGR's Terrestrial Ecology Research Group, which states as its goals, 1) to broaden knowledge of key ecological processes in Mediterranean ecosystems, 2) to develop conceptual tools and methodologies to improve the conservation and management of natural resources, and 3) to promote the transfer of research results to the management of protected areas, disseminating this information through training and advisory activities. Dr. Castro has a very extensive record of publications and presentations in the field. More information available at www.ugr.es/~rnm220/.