Wildfire effects on macroinvertebrate communities in Mediterranean streams Efectes dels incendis forestals sobre las comunitats de macroinvertebrats en rius mediterranis Iraima Verkaik ADVERTIMENT. La consulta d’aquesta tesi queda condicionada a l’acceptació de les següents condicions d'ús: La difusió d’aquesta tesi per mitjà del servei TDX (www.tesisenxarxa.net) ha estat autoritzada pels titulars dels drets de propietat intel·lectual únicament per a usos privats emmarcats en activitats d’investigació i docència. No s’autoritza la seva reproducció amb finalitats de lucre ni la seva difusió i posada a disposició des d’un lloc aliè al servei TDX. No s’autoritza la presentació del seu contingut en una finestra o marc aliè a TDX (framing). Aquesta reserva de drets afecta tant al resum de presentació de la tesi com als seus continguts. En la utilització o cita de parts de la tesi és obligat indicar el nom de la persona autora. ADVERTENCIA. 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TESI DOCTORAL Departament d‘Ecologia Programa de doctorat: Ecologia Fonamental i aplicada Bienni 2003-2005 Wildfire effects on macroinvertebrate communities in Mediterranean streams Efectes dels incendis forestals sobre las comunitats de macroinvertebrats en rius Mediterranis Memòria presentada per Iraima Verkaik Per optar al grau de Doctora per la Universitat de Barcelona Barcelona, setembre de 2010 Vist-i-plau dels directors de la tesi Dr. Narcís Prat i Fornells Dra. Maria Rieradevall i Sant Catedràtic del Dept. d‘Ecologia Professora titular d‘Ecologia Universitat de Barcelona Universitat de Barcelona The work presented in this thesis was funded by: • The Ministerio de Educación y Ciencia with scholarship BES-2005-9682, funded research stays for the 2007, 2008 and 2009 years, and projects FURIMED 1 CGL2004- 01549 and FURIMED 2 CGL2008-03388. Also contributed financially: • Universitat de Barcelona for conference assistance in 2009 • Institut de l‘Aigua for conference assistance in 2010 • Universitat de Barcelona for grant for the completion of the thesis in 2010 And received support by: • Diputació de Barcelona A los mismos 3 gatos, ahora con gatitos, y al mismo perro en el mismo chiringuito que nos sigue esperando en la playa Liniers Contents General introduction ………………………………………………………………...….9 Objectives ………………………………………………………………………...……17 Chapter 1 ……………………………………………………………………………….19 Midterm macroinvertebrate community response to different local riparian canopy affectation by a wildfire Chapter 2 …………………………………………………………………………….…43 Wildfire versus seasonal drought shaping a Mediterranean macroinvertebrate community Chapter 3 …………………………………………………………………………….…63 Effects of bushfire on structural and functional parameters of macroinvertebrate communities in Victorian creeks affected by a decade of drought Chapter 4 …………………………………………………………………………….…83 Effects of wildfire on stream macroinvertebrate assemblages in three biomes: do climate and hydrology mediate responses? General discussion ……………………………………………………………………101 Conclusions.…………………………………………………………………………..115 Resumen (en castellano)..……………………………………………………………..117 Bibliography…………………………………………………………………………..143 Appendix……….……………………………………………………………………..157 Aknowledgments……………………………………………………………………...164 General introduction Fire as a natural disturbance Wildland fire is a critical component in the terrestrial and atmospheric dynamics of our Earth system (Flannigan et al. 2009). This is not a recent story. During most of the Earth‘s history, fires have been integral to the evolution of flora and fauna, and responsive through feedback processes to changes in climatic and paleoatmospheric variations (Pausas & Keeley 2009). Burning in the last decades an average of 383 Mha annually (Schultz et al. 2008), wildfire is the natural disturbance that influences the bigger areas through out several regions (Lavorel et al. 2007). This becomes very clear when looking at the fires map from the European Space Agency (ESA), based on thermal infrared satellites images taken from July 1996 until August 2010 (Figure 1). Actually, wildfire is the key factor to understand many of the world‘s biomes distribution and fire-prone ecosystems structure and function (Bond & Keeley 2005). Fire also affects many fundamental ecosystem processes such as nutrient cycling (Turner et al. 2007), vegetation succession pathways (Turner et al. 2007; Brown & Smith 2010), and other disturbance patterns such as pest outbreaks (McCullough et al. 1998), hydrology (Shakesby & Doerr 2006), and erosion (DeBano et al. 1998; Shakesby et al. 2007). In summary, fire has been playing a significant role in the natural global carbon cycle and in regulating terrestrial ecosystems and biodiversity (Whelan 1995). At the same time, fire has been also considered the main climate-triggered hazard that can threat ecosystems and human security (Dube 2009), because biomass burning plays the major role in global environmental changes, influencing atmospheric composition, climate systems, human health, and economic activities (Schultz et al. 2008). Annual costs of fire prevention and suppression in the US have reached nearly $3 billion annually in recent years (Marlon 2009), with increases of 11% per year on average (Schoennagel et al. 2009). Fire costs damages can be very high; as an example, recent Russian fires have been calculated in $15 billion (Wikipedia 2010, http://en.wikipedia.org/wiki/2010_Russian_wildifres, retrieved September 2010). Therefore, the interest in fire and its effects has become global and interdisciplinary due 9 Introduction to influences, interactions, and feedbacks among fire, terrestrial, and atmospheric systems (Krawchuk et al. 2009). Although the physical process is theoretically simple, modern fire regimes are an ongoing focus in ecological research, nuanced by the role of humans who are changing the landscapes to be more or less flammable, and also lighten and extinguish fires (Lavorel et al. 2007; Chuvieco et al. 2008; Krawchuk et al. 2009). Black years are becoming more common, and the recent fires in Russia (2010) are again a good example with more than 300 thousand of forest hectares burned (Nature news, doi:10.1038/news.2010.404, retrieved September 2010). Moreover, these black years are more recurrent in extended areas of the world. For example, in 1994 more than 2 million hectares burned in areas such as Mediterranean climate areas, South Australia, Russia and South America (Terradas 1996); in 1997-1998 the WWF entitled this period as ―The Year the World Caught Fire‖ (Rowell & Moore 2000); in 2003 large fires burned South Eastern Australia, Western Canada, Mediterranean Europe and Southern California (Lavorel et al. 2007). All of them drew again considerable public and political attention to fire as a phenomenon through which ecological and human dynamics collide (Lavorel et al. 2007). The most recent example is the 50% rise in wheat prices partly as a consequence of the wildfires that occurred during the last summer of 2010 (BBC, http://www.bbc.co.uk/news/business-10851170, retrieved September 2010). Some models already predict a higher recurrence in fires (Westerling et al. 2006), but the future trends of global fire activity (severity and recurrence) are varied and difficult to determine owing to the complex and non-linear interactions between weather, vegetation, and people (Flannigan et al. 2009). The uncontrolled and widespread wildfires that occurred in 1997 seemed to be a consequence of extreme dry conditions brought by El Niño that year (Levine 1999). Although weather forecasts predicted 1999 as a wet year, 18 months after some of the worst forest fires in the modern era, fires started again (Rowell & Moore 2000). Thus, in general, dry years enhance the fire consequences (Westerling & Swetnam 2003), and in the future an increase in the frequency and intensity of extreme droughts is expected (Houghton et al. 1996). 10 Introduction Figure 1. Fire global map based on the ESA global detection of hot spots with twin radiometer sensors (along track scanning radiometer and Envisat‘s advanced along track scanning radiometer) from