The critical concept of scale “…the problem of relating phenomena across scales is the central problem in biology and in all of science”. Simon Levin, 1992. Why be concerned about scale? • Characterization of biogeographic pattern involves: – Scale of the phenomena – Scale of its detection through observation – Scale at which a pattern is statistically analyzed and communicated. Cartographic scale From Lewis Carroll’s Alice in Wonderland Cartographic scale Representative fractions 1:250,000 1:50,000 1:2500 Cartographic scale • Assuming that you have two maps of the same paper size, which of the two representative fractions shows the most detail and the smaller surface area? 1:50,000 1:10,000 Large versus small cartographic scale Large cartographic scale Small cartographic scale Absolute and relative scaling • Absolute scale – Distance is physical, measureable • Relative scale – Distance is not a physical distance but a more intangible construct, often similarity in some property Ecological topology • Geometric representations of ecological entities and phenomena in varying degrees of abstracted space Eulerian topologies • Connectivity between objects – Example: Networks Poincareian topologies • Field-based, geometric representation of the dynamical system states – Example: state space Operational scale • Refers to the spatial and temporal dimensions of an object or a process Operational scale • Operational scale may not be independent of our observations, and in fact may be very dependent upon them. • Operational scale can be constrained by our technology and our senses. And so these men of Indostan Disputed loud and long, Each in his own opinion Exceeding stiff and strong, Though each was partly in the right, And all were in the wrong! John Godfrey Saxe 1878 Decreasing (finer) grain, extent is constant Ecological scaling • Extent: spatial (or temporal) dimension of an object or process observed or analyzed • Grain: level of spatial (or temporal) resolution at which an object or process has been measured or observed. Increasing extent, grain is held constant Which map shows more grain? How much grain is shown can be a subjective decision. The map maker has to decide upon grain given the extent. Map generalization • Cartographers decide on the amount of generalization in a map. • Unavoidable issue related to cartographic and ecological scale • All maps have been generalized because all the detail on the surface cannot be conveyed on a paper map. Haggett’s scale coverage problem • Nature has an immense extent and a fine grain Haggett’s scale coverage problem • To make even small descriptions of it, we have to sample • Sampling requires: – Sacrificing grain for extent – Sacrificing extent for grain. Fallacies of scale • Individualistic fallacy: extrapolating to the broad scale based on observations conducted at small, local scales • Ecological fallacy: making local-scale characterizations based on broad-scale observations. • To an extent, humans are going to commit these fallacies in order to understand the world Example: local microclimates • Local buffering of climate change due to microhabitats • Cannot infer detail of local temperatures based on macroscale (large extent and coarse grain), nor can we extrapolate up from the microscale (small extent and fine grains) to the macroscale Modifiable areal unit problem • Two components – Aggregation problem – Zoning problem Hierarchy theory • Landscapes organized into spatial and temporal domains of shaping processes • System of vertical interconnections • Higher levels set the context for the lower levels Hierarchy theory Example: how hierarchical theory informs the assembly of communities Example: hierarchy theory and extinction filters Non-hierarchical controls also relevant Examples: Tilt, insolation, climate, photosynthesis: hierarchical Invasive species, dispersal, natural selection: nonhierarchical Cartesian scale • Hierarchy theory exemplifies a Cartesian scaling of the world • Scales are imposed • Not necessarily “true” scales • Often tied to XYZ coordinate system • Space as a container Constructivist scale • Scale is produced by organisms • There is no fixed and unchanging hierarchy of scales imposed from above. Constructivist scale • Boundaries in space and time shaped by organisms not just our observation of them. • Extents and grain are also a function of the organisms rather than our (human) sensing alone The epistemological mode of scale • Epistemology: strategy of belief that informs us of what we can and cannot know • Epistemological mode generates information useful for ascertaining predictability • Organisms scale the world to promote predictability or the appearance thereof • Our senses, technology, and beliefs constrain scales of our perception and consequently what we can claim to know. The ontological mode of scale • Ontology: what exists, the categories by which we label and define the world • Other organisms also scale and act upon their inferences of predictability • Borders and boundaries are not universally shared • Introduces unpredictability – scale mismatches The biological mode of scale • The interaction of the epistemological and ontological moments of scale in organisms • Epistemological scaling needed for predictability • Ontological scaling produces unpredictability • Microorganisms to complex organisms scale the world to generate predictability – they infer and act upon the extent (boundaries) and grain (or quality) of phenomena perceived in time and space • Unpredictability arises from environmental change and other organisms that also infer, act upon, and modify their scaling of the environment The biological mode of scale • Basis for adaptation and evolution • How have the extents and grains detected by our senses and technology been intertwined with our evolution? Rules for ecological scale for humans 1. Patterns are dependent upon the scale of observation 2. The important explanatory variables change with scale. 3. Statistical relationships may change as scale changes. 4. Patterns are generated by processes acting over various temporal and spatial scales. Patterns are dependent upon the scale of observation Patterns are dependent upon the scale of observation The important explanatory variables change with scale. The important explanatory variables change with scale. Patterns are dependent upon the scale of observation The important explanatory variables change with scale. Patterns are dependent upon the scale of observation Statistical relationships may change as scale changes. Rules for ecological scale 5. Scale can be used to justify or refute certain management practices and ideas about nature Example: Successional response to clearcut logging • Grain and extent of post-logging sampling determine criteria for judging response – Resurvey over large extent and fine grain: criteria for recovery not likely to be met – Resurvey over small extent and coarse grain: criteria for recovery easier to meet Rules for ecological scale 6. The scales experienced by an organism define what it sees and responds to. For example, what might constitute a patchy resource to an insect, could be perceived by a larger vertebrate as homogeneous. How to work with scale • There is no single correct scale or level at which to describe a system. • This does not mean that all scales serve equally well or that there are not rules or guidelines. How to work with scale • Be aware of the different types of scaling • Don’t be too anthropocentric • Employ sampling designs and methods that are sensitive to multiple scales – – – – Nested observations Power laws Fractals Networks Nested observations Sp - species Power laws • Summarize how relationships change with changes in extent • Often expressed on a log-log plot. • Y = constant (X)n • Similar slopes are thought to have similar structuring processes (n = slope) • Examples • • Species-area relationships (left): S=cAz Animal metabolic rates and body mass (next slide) • However, a valid criticism of power laws is that they often lack an explanatory process Metabolic rate and body mass scales according to this power law: Metabolic rate = constant (mass).75 (Metabolic rate) (Mass) Fractals • A fractal pattern appears the same across all scales. It is scale invariant. • Fractal patterns can be expressed as a power law with a particular range of slope values. • Power laws do not necessarily imply fractal structure. Networks • Can represent relationships at a variety of scales at once. • Eulerian topology • Structural properties of networks provide means of understanding how they work – Nodes and links – Degree centrality and betweenness – Weak versus strong links – Directional versus nondirectional graphs Random versus scale free networks can reflect different modes of assembly and activity. These may also have implications for their geographic distribution Spatial autocorrelation • A method to summarize how patterns change with scale. • Based on Tobler’s Law: near things are more likely to be similar than things at a greater distance apart