Bioe 147/247 Lecture Notes Community Ecology: Basic Concepts and Approaches Community Ecology = The Ecology of Biodiversity Community ecology and biodiversity deal with difficult issues: Many species: different evolutionary histories & different roles Community patterns and processes vary in details Type of system (aquatic v terrestrial; altitude; latitude, etc.) Stage of community development (e.g., time since disturbances) Adjacent/nearby communities: Regional vs Local effects Abiotic factors (e.g., climate (always changes); nutrients, disturbances) BUT: there are many similarities of processes & principles across different communities, and I’ll stress them in this course. PROBLEM: What is a community? Many ways to define community reflect different approaches & types of questions Morin & most other texts: define community as a collection of spp coexisting together in space & time. This is: very weak, too general and misleading. ignores what species do. This definition is also confusing and ambiguous because it leaves open these questions: 1. What is coexistence? - Just being there? - Interaction? - How long? - Species evolve – so what units are coexisting? Do spp do more than just ‘coexist’? 2. What spatial scales? - grassland plants & how many neighbors? Forests? Open ocean? - Some species only use the same space infrequently: don’t co-occur most of the time, but interact in important ways (e.g., plant/rabbit/coyote) DRAW Species can affect others even if they don’t share the same space most of the time. 1 3. What temporal (time) scales? - ecological events among interacting spp can differ. - trees, seeds, insect herbivores DRAW Spp can affect others even if their time frames of existence are very different. There are 3 different approaches: (see Table 1 – pgs 3-4) to studying communities that reflect different ways to address those problems in defining coexistence and scale. These approaches also address different questions. For this class: we use all approaches as we address different aspects of communities. Definition: For this class, we will use the following definition (after TFH Allen) to avoid some of the usual problems. An ecological community is a) an assemblage of species that occurs together because of interactions that have led to ecological or evolutionary accommodations among the species; and b) an assemblage of species that may also reflect regional processes (e.g., dispersal, disturbance and evolution beyond a local community). Main words: interactions, accommodation, regional Why accommodation? (presence affects others) Why evolution? e.g. Competition NOW Exclusion of weak competitors Changes in competitive relationships (‘The Ghost of Competition Past’) PAST Extinctions (reduce# of spp) Character displacement interactions use different resources (note: this can be very fast- in only a few generations) Why regional? Dispersal different effects? not evolutionary? 2 With this new definition, there are no problems viewing the coyote-rabbit-plant interactions as in the same community they form a chain of interactions even though they use both space and time differently. Major Themes of Community Ecology: - Mechanisms/processes that determine membership (especially interactions) - Patterns: arising from mechanisms 3 TABLE 1: NOTE: I’ve circled the important features of each approach that you should know and think about. Assemblage Approach Functional Group Approach Interactions Approach Groups: Main Qs relate to: Combines elements of assemblage & Interactions among species Broad groups: species approaches to: Taxa (birds, butterflies…) Trophic level (plants, Uses: subsets of spp (fn gps = guilds) based carnivores…) on roles in communities or by morphology, OR physiology, etc. Particular taxa (eg, hawks) w/in groups e.g., N-fixing plants, encrusting algae, sapsucking herbivores, parasites on flies…. Focus Compare taxa (groups of spp) among samples Assess - Group traits: S, biomass… - Species ID &/or abundance - Compare different fn gps (properties, interactions, responses, etc.) - Responses and roles of the fn group(s) and NOT the component species Mechanisms (= processes) that allow membership in local community - Species’ responses to other spp (over time &/or space) General methods Field (tactics) Analyses Observations (mostly) Experiments (some) between groups Statistics - quantify differences & similarities of samples: classification, ordination; some tests of hypotheses Observations (some) Experiments (some) between fn gps Statistics a) classification/ordination methods to help id the fn groups and compare samples/communities; b) test hypotheses about differences betw/ fn gps. Experiments (mostly) Observations (some) among species Statistics - test hypotheses re interactions and processes 4 Strengths - ID broad patterns between - Can ID either patterns and mechanisms different communities or samples (depending on the specifics of a study) - May ID some mechanisms (if do expts or id key features of the environment that result in particular communities patterns) What do we learn? Limitations - Emphasizes that spp play active role in community - Spp don’t occur in rigidly defined groups - Spp may belong to several communities - Group (taxon) patterns vary in space & time - At any time/place – spp present are subsets of potential members many more spp can live there than do. - Properties of the group as a whole can swamp those of individual/particular species. - Largely descriptive - Groups can be defined by different functions and this could lead to different groupings. - All spp are assumed equal - Ignores what spp do - Cannot id mechanisms w/in community - Can ID mechanisms of membership in community (not just pattern) e.g.: Properties of groups (such as biomass) may be constant even though the particular species may vary considerably [see reading by Hay about species as ‘noise’ for wk 2] The composition of groups may change if different criteria are used (e.g., defining spp according to resource use yields a different grouping than if use physiological criteria). Which is right??? Both, but that leads to some complications. - membership in a community may be active, & due to: a) population processes (incl some behaviors) of species that affect b) interactions among species, & c) evolution in response to other species. OR smts: - d) dispersal - What is an interaction? - How strong an interaction is needed to affect other spp or community properties? - Reductionist Detailed expts on only a few spp (usu the most common) ‘Cartoon” of communities 5 (See - pg 11 of supplement). In addition: there is often a big difference between ‘textbook’ science and ‘frontier’ science. In community ecology, I do as much ‘frontier’ science as possible – which means that ideas are changing/evolving as we get more information. FOR this class: we use examples from terrestrial, marine and freshwater communities. I’ve selected the best examples to make each point, regardless of the system. FINALLY: I have a political motive here because everyone needs to be concerned about biodiversity and because biodiversity and community ecology are closely linked. I want to make sure that you (and I) work hard so that you are scientifically literate re community ecology. Some comments [from an Op-Ed piece by Brian Greene in the New York Times (6/1/08)] about science in your life and about science and social responsibility: - It is crucial to cultivate a “general public that can engage with scientific issues; there’s simply no other way that as a society we will be prepared to make informed decisions on a range of issues that will shape the future.” - Too often “our educational system fails to teach science in a way that allows students to integrate it into their lives”. - “…science can play” a powerful role “in giving life context and meaning”. - “Science is a way of life. Science is a perspective. Science is the process that takes us from confusion to understanding in a manner that’s precise, predictive and reliable — a transformation,” - “…science is still widely viewed as merely a subject one studies in the classroom or an isolated body of largely esoteric knowledge that sometimes shows up in the “real” world in the form of technological or medical advances.” Or else people “feel that the science itself is just not relevant to their lives.” Knowing about science is a social responsibility. This course deals with material critical to major current issues in your lives and our society, especially: Biodiversity (by definition of community ecology) Conservation Climate Change We don’t discuss solutions, but you need to know the deep issues behind the problems. 6 Basic Concepts in Community Ecology & about Biodiversity Basic questions in Community Ecology/ Biological Diversity (mechanisms membership of species in a community): 1. Community Boundaries 2. Do communities recur in space and/or time? How similar are they? 3. Why do different communities have different numbers of species? 4. What are the patterns? What causes them? 5. What is the role of history vs. current interactions? 6. What is the role of regional vs. local processes? 7. What are ‘emergent’ properties of communities (e.g., stability, productivity, succession) and how are they affected by species diversity, or by particular species? Major Themes: 1. Species diversity (richness) 2. Interactions 3. Change 4. Scale: space, time (implicit) Tactics – how the course is structured: 1. Community Patterns = assemblages, S, similarities, diversity, food webs, roles 2. Short-term Dynamics = multi-species interactions 3. Emergent Properties (linking structure dynamics): stability, coevolution 4. Long-term Dynamics = change, evolutionary patterns, history (succession, evolution, climate change, site change [e.g., disturbances], future changes Finding Community-level Generalizations is hard: o Spp. have different evolutionary histories o Spp. have different ecological roles o Community patterns and processes may vary with - Type of system (terrestrial or aquatic; abiotic properties (e.g. altitude, climate)) - Stage of community development (time since disturbance). - Nearby communities regional vs local impacts In addition: there is often a big difference between ‘textbook’ science and ‘frontier’ science. In community ecology, I do as much ‘frontier’ science as possible – which means that ideas are changing/evolving as we get more information. 7