Stream Ecology (NR 280) Topic 10 – Lotic Communities Review of community elements Large-scale influences on species diversity What controls community structure? Do species prefer one niche over another? How does community structure change over time? Community structure and ecosystem function Using community structure for management Building blocks of community structure in stream communities • Autotrophic species – Algae – Bryophytes – Macrophytes • Micro-heterotrophic species – Taxonomic groups (e.g., Pseudomonads) – Functional biogeochemical groups (e.g. denitrifiers, sulfate reducers) • Benthic macroinvertebrate species – Taxonomic groups (e.g., EPT) – Functional feeding groups (e.g., grazer, shredder, etc.) • Fish species – Taxonomic groups (e.g. Cyprinids, Salmonids) – Fish guilds (e.g., bottom feeder, algivore, piscivore) • Stream-reliant species – Salamanders, birds, bears Interactions within community structure that define stream communities • Food and Feeder – Primary production and grazing/herbivory – Secondary production and consumption – Predation and cannibalism • Functional relationships – Competition – Mutualism/Commensalism – Special species interactions Expressing community structure Connectance food web - Broadstone Stream, UK Schmid-Araya et al. (2002) in Allan and Castillo (2007) Expressing Strength of Connection Example: Energy Flux Food Web Ogeechee River, Georgia Benke and Wallace (1997) in Allan and Castillo (2007) Interesting questions to consider • What is a community? • Why don’t we see the same species everywhere? • What factors influence the structure of biological communities in stream ecosystems? • What is the influence of community structure on ecosystem function? Stream Ecology (NR 280) Topic 10 – Lotic Communities Review of community elements Large-scale influences on species diversity What controls community structure? Do species prefer one niche over another? How does community structure change over time? Community structure and ecosystem function Using community structure for management Species Area Curves Two different senses of this concept • Sense 1: Diversity = f(area available) – A metric of the environment – A indicator of regional diversity • Sense 2: Richness = f(area sampled) – A metric of sampling effort – A indicator of local diversity Species Area Relationships Larger areas typically support more species 49 coastal streams on US East Coast to Gulf of Mexico. Note log-log axes and linear form. Sepkowski and Rex (1974) in Allan and Castillo (2007) Remember that this relationship has the familiar log-linear form S = c Az S = number of species c = constant A = area z = scaling factor (constant) Log(S) = Log(cAz) Log(S) = Log(c) + Log(Az) Log(S) = Log(c) + z(Log(A)) Y = a + bX Power function = Log-Linear function Relationship between basin area and fish diversity in world rivers Mekong River Commission Species Area Curves Two different senses of this concept • Sense 1: Diversity = f(area available) – A metric of the environment – A indicator of regional diversity • Sense 2: Richness = f(area sampled) – A metric of sampling effort – A indicator of local richness Local species area (sampling effort) The species area relationship is shown from the rate at which species are added to collections as sampling effort (shown by number samples) increases. In this example, samples of macroinvertebrates have been collected from four habitats (2 pools, 2 riffles) in a Hong Kong stream. Note that the species accumulation curve does not level off in Riffle #1 because new, rare species are encountered even after 30 samples have been taken. Mekong River Commission Species by abundance Are all species equally abundant? Broadstone River United Kingdom 24 years of sampling Woodward et al. (2002) in Allan and Castillo (2007) Special Species Interactions Unionid mussels and fish Watters (1992) Special Species Interactions Unionid mussels and fish Unionid mussels require specific fish hosts to transport the larval form of the mussel (glochidia). Watters (1992) Stream Ecology (NR 280) Topic 10 – Lotic Communities Review of community elements Large-scale influences on species diversity What controls community structure? Do species prefer one niche over another? How does community structure change over time? Community structure and ecosystem function Using community structure for management What controls community structure? The deterministic view • Communities are non-random assemblages of species with predictable and repeatable structural patterns • Communities are relatively stable in the face of “ordinary” environmental stresses • Community structure is the outcome of interactions between habitat characteristics and species preferences What controls community structure? The non-deterministic view • The specific structure of a community is dependent on a random sample of species from the regional pool that are able to survive and reproduce in the local environment • Hubbel’s (2001) “Neutral model”: Species are interchangeable. The habitat offers a template (deterministic) that is then randomly occupied by species (non-deterministic) Key theories about stream community structure These are not mutually exclusive • Habitat Template influences: physical characteristics of the habitat determine what species can exist in a place; diverse habitats have diverse community structure • Disturbance influences: environmental disturbance (e.g. flow, temperature) favors some species and does not favor others • Niche influences: structure is determined by interplay of biotic interactions (e.g. predation, herbivory, competition) and abiotic forces (habitat, disturbance) Some general observations • Community structure is often persistent over years • The same species are often dominant at a location, year after year But… • High temporal variation in key environmental variables may mask persistence and dominance • Persistence and dominance are indicative of deterministic factors but don’t explain them Stream Ecology (NR 280) Topic 10 – Lotic Communities Review of community elements Large-scale influences on species diversity What controls community structure? Do species prefer one niche over another? How does community structure change over time? Community structure and ecosystem function Using community structure for management Why should individual species prefer specific habitats? The concept of species traits • Species have particular traits that may impose specific habitat requirements • Habitats offer specific characteristics that may favor particular species Lamouroux et al. (2004) Example species traits & habitat characteristics 16 of 60 traits used in this study Lamouroux et al. (2004) What Lamouroux et al. concluded • Strong relationship between species traits and microhabitat characteristics • Consistent relationships at the reach scale • Inconsistent relationships at the basin scale (n=2) Poff (1997), also Fig 1.5 Allan and Castillo (2007) General Observation Local biological diversity is the outcome of regional species richness interacting with local environmental conditions and historical events to define the subset of species that exist at a given time in a given stream location. Paraphrased from Allan and Castillo (2007), p. 230 Regional Definition/Local Expression Fixed Environmental Template Filters; e.g. Geology Topography Insolation Regional Species “Reservoir” Changing Environmental Template Local Habitat Filters; e.g. Runoff Climate Glaciation Local Community Local scale History (presses and pulses) State (conditions) Regional scale Stream Ecology (NR 280) Topic 10 – Lotic Communities Review of community elements Large-scale influences on species diversity What controls community structure? Do species prefer one niche over another? How does community structure change over time? Community structure and ecosystem function Using community structure for management Streams have different flow regimes Based on 78 USGS gauging stations High Low How intermittent is the flow? Not at all Harsh Intermittent Abiotic Intermittent Runoff Abiotic Perennial Flashy Low flow predictability Abiotic Snow and Rain Low flow predictability Seasonally biotic Perennial runoff Low flow predictability Abiotic Winter rain Low flow predictability Seasonally biotic Mesic groundwater High flow predictability Biotic Snowmelt High flow predictability Seasonally biotic Flood Frequency High Low Intermittent Flashy Abiotic Low High Flood Predictability Modified from Poff and Ward (1989) in Allan and Castillo (2007) Flow regime, disturbance, & community structure Disturbance causes short-term change Insect abundance in a stream in the Andean foothills after >25mm storm events Flecker and Feifarek (1994) in Allan and Castillo (2007) Recovery following flash flood disturbance Sycamore Creek, Arizona Autotrophs Benthic Macroinvertebrates Fisher et al. (1982) in Allan and Castillo (2007) How communities change over time Long-term factors • Dispersal – movement of species in space – Immigration: species moving to a location – Emigration: species moving from a location • Speciation – development of species in time • Biogeography – suitability of species to exist in a particular space at a particular time Presses, pulses, and tipping points • Press – A steady application of perturbing force – Climate change, development – A measure of resistance to change • Pulse – A sudden application of a perturbing force – An accidental spill, Hurricane Irene – A measure of resilience to change • Presses and pulses may interact – Non-linear responses, “surprises” – Tipping points – permanent changes to a new state Stream Ecology (NR 280) Topic 10 – Lotic Communities Review of community elements Large-scale influences on species diversity What controls community structure? Do species prefer one niche over another? How does community structure change over time? Community structure and ecosystem function Using community structure for management Interspecific facilitation Filtration of suspended particles in artificial stream • 3 filter-feeding larval caddis fly species • Feeding alone (open) or with other 2 species (hatched) • Dependent variable is per capita consumption of suspended matter Cardinale et al. (2002) in Allan and Castillo (2007) Interspecific Facilitation Impacts on decomposition of leaves High dominance Low dominance Low evenness High evenness 36 streams France Sweden Low dominance ○ ● High dominance □ ■ Damgles and Malmqvist (2004) in Allan and Castillo (2007) Removing one species alters ecosystem processing Atlas of the freshwater fish of Venezuela Taylor et al. (2006) Difference -fish -fish Data Experiment results Benthic organic carbon Carbon flux Heterotrophic respiration (only) Gross primary production P:R ratio (actually GPP/CR) Net ecosystem metabolism Carbon turnover length Taylor et al. (2006) See comments panel for legend Human impact: smaller fish (smaller net sizes, not shown) Stream Ecology (NR 280) Topic 10 – Lotic Communities Review of community elements Large-scale influences on species diversity What controls community structure? Do species prefer one niche over another? How does community structure change over time? Community structure and ecosystem function Using community structure for management Common Benthic Macroinvertebrates Moderately Tolerant Sensitive (Intolerant) • • • • • • • • Stoneflies Water Penny Beetles Mayflies Dobsonflies Alderflies Snipeflies Mussels Riffle Beetles • • • • • • • • Damselflies Dragonflies Crayfish Amphipods Blackflies Caddisflies Isopods Craneflies Tolerant (Insensitive) • • • • Midgeflies Worms Leeches Pouch Snails http://www.epa.gov/bioindicators/html/benthosclean.html Benthic Macroinvertebrates The “Good” EPT Taxa The “Poor” taxa http://www.epa.gov/bioindicators/html/benthosclean.html Simple guide (North Carolina) Macroinvertebrate Index of Biotic Condition • • • • • • Taxonomic richness Taxonomic composition Taxonomic diversity Feeding groups Habitats Pollution tolerance EPA Wadeable Stream Assessment (2007) Index of Biotic Integrity Karr (1981) and Karr et al. (1986) from EPA-260-R-08-016 (2008) Response of Sensitive EPT Biota High-gradient stream reaches Non-linear response at ~5% TIA No significant response at TIA > 10% (N = 6; p = 0.157) % Cobble substrate explained 11% of EPT variance Watershed TIA Fitzgerald et al. (2012) Impacts of Impervious Area Center for Watershed Protection (2003) as noted.