13th Century model of rabbit population growth:

1. 13th Century model of rabbit population growth: based on Fibonacci series (1, 1, 2, 3, 5, 8, 13, ….) Malthus (1798) … Nature has scattered the seeds of life abroad with the most profuse and liberal hand. She has been comparatively sparing in the room and nourishment necessary to rear them. The germs of existence contained in this spot of earth, with ample food, and ample room to expand in would fill millions of worlds in the course of a few thousand years…. Given unlimited resources, size of population increases as geometric progression(1, 2, 4, 8, 16, ….) Verhulst-Pearl logistic equation (1839) model of population growth in a limited environment

2. Annual Calcium Budget for an Aggrading Forested Ecosystem at Hubbard Brook (Likens et al. 1977) NORTHERN HARDWOOD FOREST ECOSYSTEM ABOVE GROUND LIVING BIOMASS BOUND Ca 383 (5.4) Inorganic fraction 2.2 LITTER FALL 40.7 THROUGHFALL AND STEMFLOW 6.7 TRANSLOCATION JJJJJJJJJ JJJJJJJJJJJJJJJJJJ -5.1 INPUT BELOW GROUND LIVING BIOMASS BOUND Ca 101 (2.7) BULK PRECIPITATION 2.2 UPTAKE 62.2 FOREST FLOOR BOUND Ca 370 (1.4) BIOSPHERE ROOT LITTER 3.2 HYDROLOGIC EXPORT 13.9 ROOT EXUDATES 3.5 Inorganic fraction 3.5 SOIL AVAILABLE Ca 510 Organic fraction <0.1 Inorganic fraction (particulate) 0.2 NET MINERALIZATION 42.4 OUTPUT MINERAL SOIL BOUND Ca 9600 ROCK 64,600 WEATHERING 21.1 dissolved organic fraction 13.7

3. A conceptual model is a mental picture of how something works. We have a conceptual model of a car that allows us to drive by relating certain actions (e.g. pressing the brake pedal) to certain results (e.g. the car stops). We don’t have to understand automotive engineering for our driving model to work. But if we need to repair the engine, a different model would be required.

4. Why Aren’t Conceptual Models Routinely Used to Develop Ecological Monitoring? Cynicism regarding utility of modeling Incomplete understanding of ecosystem function Confusion over modeling objectives Prototype Confessions . . . . . We did not initiate the Prairie Cluster LTEM program with formal planning process -- convening panels of experts. We are reviewing monitoring components within context of ecosystem models rather late in the design phase.

5. Sparse and Infrequent Observations Observational Errors Incorrect Interpretation Theoretical Misunderstanding Management Decisions Oversimplified Models Further Refinement of Unimportant Details Computer Models CONTROVERSY Unrealistic Assumptions CONFUSION Further Misunderstanding Crude Diagnostic Tools Coincidental Agreement Between Theory and Observations PUBLICATION Cynic’s View of the Interface Between Ecological Research and Management (Hobbs, 1998)

6. Tactical Models versus Strategic Models (May 1973) attempt to measure all relevant factors way of formalizing generalizations and determine how they interact about the ecological system of interest “a purposeful representation of reality” (Starfield et al. 1994) • Common Misconceptions(Starfield et al. 1997) • A model cannot be built with incomplete understanding.Managers make • decisions with incomplete information all the time! This should be an added • incentive for model-building as a statement of current best understanding. • A model must be as detailed and realistic as possible. • If models are constructed as ‘purposeful representations of reality’, then design • the leanest model possible. Identify the variables that make the system behave and • join them in the most simple of formal structures.

7. Modeling Confusion? • Conceptual Models of Indicator Selection Process • Conceptual Ecosystem Models • Small, Focused Models -- Conceptual or predictive models of populations • or communities • Holistic Program Models -- Conceptual models of how monitoring • information will feed back into • decision-making process

8. Conceptual Models of Indicator Selection Process • Conceptual Ecosystem Models

9. Conceptual Ecological Models of the Major Wetland Physiographic Regions in South Florida Comprehensive Everglades Restoration Project Team and the Science Coordination Team of the South Florida Ecosystem Restoration Working Group Lake Okeechobee Caloosaatchee Estuary St. Lucie Estuary & Indian River Lagoon Everglade Ridge and Slough Big Cypress Basin Southern Everglades Marl Prairies Southern Shark Slough Mangrove Estuary Transition Florida Bay Mangrove

10. Exotic Plants Sea Level Rise Reduced Freshwater Flow Volume & Duration Exotic Fish Altered Nutrient Mixing & Estuarine Productivity Altered Salinity Gradient Coastal Embankment Erosion Exceeding Accretion Altered Hydroperiod & Drying Patterns Invasion of Schinus & Colubrina Altered Mangrove Production Community Structure & Organic Sediment Accretion Invasion of Mayan Cichlid Estuarine Fish Communities & Fisheries Estuarine Geomorphology Mangrove Forests & Plant Communities Woodstork & Roseate Spoonbill Nesting Colonies Estuarine Crocodilian Populations Conceptual Model of Mangrove Estuary Transition

11. Conceptual Models of Indicator Selection Process • Conceptual Ecosystem Models • Small, Focused Models-- Conceptual or predictive models of populations or communities

12. spatial variability at very small scales geology shallow soils, prone to drought and frost heave climate, climate change, elevated CO2 temporal variability at multiple scales woody overstory development resource/nutrient availability woody species removal vegetation structure & composition habitat quality highly variable in time & space wildfire, prescribed fire spatial & temporal variability exotic species establishment increased edge effect differential germination, survival & reproduction reduced recolonization by native species habitat fragmentation Conceptual model of influences on Missouri bladderpod habitat quality Prairie Cluster LTEM Program

13. Conceptual model of influences on population dynamics of Missouri bladderpod climate, climate change, elevated CO2 autumn weather (precipitation & temperature) spring weather (precipitation, duration of flowering period) seed bank persistence fungal growth wildfire, prescribed fire pollinator activity seed bank reproduction vegetation structure & composition mature plants woody species removal germination germinated seedlings growth & survival exotic species establishment resource/nutrient availability habitat fragmentation wildlife/bird activity soil disturbance geology variable soil depth winter & spring weather (freeze-thaw,heavy rainfall) cultural use (trampling)

14. Conceptual Models of Indicator Selection Process • Conceptual Ecosystem Models • Small, Focused Models -- Conceptual or predictive models of populations or communities • Holistic Program Models -- Conceptual models of how monitoring • information will feed back into • decision-making process

15. Threats Resource Actions How is external land use Is the proximity or size of Landscape changing? nearby prairie remnants How is prescribed fire Are land-use changes changing? affecting prairie plant affecting prairie remnants? communities? Do prairie streams support diverse macroinvertebrate Are restoration Is the water quality of communities? methods working? prairie streams declining? Community Do small prairie remnants Is rare species habitat support diverse native plant Where are are invasive protection & communities? exotics distributed within restoration working? and adjacent to the Do small prairie remnants park? support diverse butterfly Are exotic control and bird communities? efforts effective? Are rare species re- Population colonization sources Are rare species populations disappearing? stable? Holistic program model for the Prairie Cluster LTEM Program Are prairie remnants sustainable within small parks? Indicators of Ecosystem Health

16. Monitoring Products Management Feedback Monitoring Effort Synthesis Adjacent Land Use External land use maps Distribution maps & population size estimates of rare species; Population models of federally endangered species How are changes in land use impacting the prairie? Rare Plant What areas harbor the highest diversity? What are the high risk habitats? Trends in plant community diversity, structure & composition; Vegetation maps How is the prairie changing? Is the prairie healthy? Plant Community Is the prairie threatened by exotic invasion? Distribution maps of invasive exotics Exotic Species How are management practices influencing the prairie? Are exotic control methods effective? Management History How is prescribed fire changing the prairie?

17. Conceptual models are useful throughout • the monitoring process: • formalize our current understanding of the context and scope of the natural processes and anthropogenic stressors affecting ecological integrity • help expand our consideration across traditional discipline boundaries • Most importantly, clear, simple models facilitate communication between: • scientists from different disciplines • researchers and managers • managers and the public

18. Conceptual Models of Indicator Selection Process • Conceptual Ecosystem Models • Small, Focused Models -- Conceptual or predictive models of populations • or communities • Holistic Program Models -- Conceptual models of how monitoring • information will feed back into • decision-making process

19. Why Do We Need Conceptual Ecosystem Models? Despite the complexity of ecosystems and the limited knowledge of their functions, to begin monitoring, we must first simplify our view of the system. The usual method has been to take a species-centric approach, focusing on a few high-profile species; that is those of economic, social, or legal interest. Because of the current wide (and justified) interest in all components of biological diversity, however, the species-centric approach is no longer sufficient. This wide interest creates a conundrum; we acknowledge the need to simplify our view of ecosystems to begin the process of monitoring, and at the same time we recognize that monitoring needs to be broadened beyond its usual focus to consider additional ecosystem components. Noon et al. 1999

20. Aspects to Consider as Conceptual Models are Developed from Barber (1994) 1. Identify the structural components of the resource, interactions between components, inputs and outputs to surrounding resources, and important factors and stressors that determine the resource’s ecological operation and sustainability. 2. Consider the temporal and spatial dynamics of the resource at multiple scales because information from different scales can result in different conclusions about resource condition. 3. Identify how major stressors of resource are expected to impact its structure and function

22. Physical impact Mycorrhizae Grazers Invertebrates Soil N storage Watershed and landscape patterns Grazer selectivity and grazing patterns Plant community structure Plant growth & demography Fire Direct Effects Insects Birds Local extirpation emigration and immigration Resource Availability Mammals Drought Standing dead & litter Conceptual model of core abiotic and biotic relationships within terrestrial prairie ecosystems. Modified from Hartnett and Fay (1998), the model has been adopted by the Prairie Cluster LTEM Program.

24. Conceptual model of core abiotic and biotic relationships within terrestrial prairie ecosystems, including anthropogenic stressors (in red) affecting Prairie Cluster parks. Modified from Hartnett and Fay 1998 Physical impact Exotic Plant Invasion Mycorrhizae Grazers, Cattle Invertebrates Watershed and landscape patterns Grazer selectivity and grazing patterns Plant community structure Plant growth & demography Prescribed Fire Direct Effects Insects Birds Local extirpation emigration and immigration Resource Availability Mammals Drought Cultural use Standing dead & litter Fragmentation

25. Monitoring implications from terrestrial prairie model

26. 2. Consider the temporal and spatial dynamics of the resource at multiple scales because information from different scales can result in different conclusions about resource condition. 3. Identify how major stressors of resource are expected to impact its structure and function Aspects to Consider as Conceptual Models are Developed from Barber (1994) 1. Identify the structural components of the resource, interactions between components, inputs and outputs to surrounding resources, and important factors and stressors that determine the resource’s ecological operation and sustainability.

27. The scale of resolution chosen by ecologists is perhaps the most important decision in the research program, because it largely predetermines the questions, the procedures, the observations, and the results. ….. Many ecologists…. focus on their small scale questions amenable to experimental tests and remain oblivious to the larger scale processes which may account for the patterns they study. P.D.Dayton and M.J. Tegner (1984) Most environmental problems are driven by mismatches of scale between human responsibility and natural interactions. Lee (1993)

28. Global Global Weather Systems Carbon Dioxide Variations Atmospheric Composition Origin of Earth and Life Climate 104 Glacial Periods Plate Tectonics El Niño Ocean Circulation Mantle Convection Upper Ocean Mixing 103 Synoptic Weather Systems Soil Development SPATIAL SCALE (km2) Extinction Events Soil Moisture Variations 102 Earthquake Cycle Soil Erosion Metallogenesis Seasonal Vegetation Cycles Volcanic Eruptions 101 Atmospheric Convection Nutrient Cycles 100 Atmospheric Turbulence Local Minute Day 100 yr 102 yr 104 yr 106 yr 109 yr Second TEMPORAL SCALE Spatial and Temporal Characteristics of Different Earth System Processes(NASA, 1988)

29. An Assessment of the Spatial and Temporal Scales of Natural Disturbances in an Arctic Tundra Ecosystem(Walker, 1991) Climate Fluctuations Associated with Glaciations Climate Fluctuations Associated with Glaciations Continental Drift and Uplift of Brooks Range 6 Climate Fluctuations During the Holocene Growth and Erosion of Ice Wedges and Erosion of Ice Wedges 4 EVENT FREQUENCY (log of years) Tundra Fires Megascale 2 Annual Fluvial Erosion and Deposition Eolian Deposition Major Storms and Storm Surges Snowbank Formation and Melting Animal Disturbances Oil Seeps 0 Daily Freeze- Thaw Cycle Thaw Cycle Microsite Mesocite Macrosite Microregion Mesoregion -2 -4 -2 0 2 4 6 8 10 12 SPATIAL SCALE (log of area in m2)

31. Incorporating Multiple Points of View Allen and Hoekstra (1992) stress that ecology is in many ways a ‘soft-system’ science, one in which point of view (ecological level of inquiry, temporal/spatial scale) is the very substance of the discourse. They suggest there are enough decision points in an ecological investigation (or in the design of a monitoring program) to require some formalization of decision -making.

32. Checkland’s “Soft-systems Methodology”(Allen and Hoekstra 1992) • 1) Recognize that there is a problem, a real mess • troubled feeling an ecosystem, community or population ecologist may have that some other sort of specialist could better solve the problem at hand • trying to manage water, vegetation and wildlife in a unit of particular size but realizing the temporal or spatial scales don’t mesh with natural process scales • 2)Actively generate as many points of view for the system as possible • -- ‘painting the rich picture’ • community ecologist consider physiological aspects of the problem, population biologist to consider nutrient cycling, etc. • 3) Find the root definitions -- develop abstractions that restrict the rich picture in hopes of finding solution. • (key system attributes will change as scale of the system and point of view (ecological discipline) is altered • 4) Build the model

33. Why Do We Need Conceptual Models? 1) Ecosystems (communities, populations) are messy; our ability to provide early warning of resource decline is uncertain. We need a road map. 2) Long-term monitoring is an iterative process (i.e. we may not get it right the first time); modeling will help ensure that mistakes are instructive and not repeated. 3) A balanced monitoring program should consider multiple spatial/temporal scales and integrate monitoring across ecological disciplines. Models serve as heuristic devices to foster better communication and clarify scaling issues. 4) A balanced monitoring program should address short-term management issues and long-term ecological integrity. Clear models serve as heuristic devices to foster better communication between managers and scientists, and between managers and the general public.