Key Words ecological boundary, ecotone, edge effect, effective area model, core area model, habitat fragmentation ■ Abstract Edge effects have been studied for decades because they are a key component to understanding how landscape structure influences habitat quality. However, making sense of the diverse patterns and extensive variability reported in the literature has been difficult because there has been no unifying conceptual framework to guide research. In this review, we identify four fundamental mechanisms that cause edge responses: ecological flows, access to spatially separated resources, resource mapping, and species interactions. We present a conceptual framework that identifies the pathways through which these four mechanisms can influence distributions, ultimately leading to new ecological communities near habitat edges. Next, we examine a predictive model of edge responses and show how it can explain much of the variation reported in the literature. Using this model, we show that, when observed, edge responses are largely predictable and consistent. When edge responses are variable for the same species at the same edge type, observed responses are rarely in opposite directions. We then show how remaining variability may be understood within our conceptual frameworks. Finally, we suggest that, despite all the research in this area, the development of tools to extrapolate edge responses to landscapes has been slow, restricting our ability to use this information for conservation and management.

Summary1. Recent advances in occupancy estimation that adjust for imperfect detection have provided substantial improvements over traditional approaches and are receiving considerable use in applied ecology. To estimate and adjust for detectability, occupancy modelling requires multiple surveys at a site and requires the assumption of 'closure' between surveys, i.e. no changes in occupancy between surveys. Violations of this assumption could bias parameter estimates; however, little work has assessed model sensitivity to violations of this assumption or how commonly such violations occur in nature. 2. We apply a modelling procedure that can test for closure to two avian point-count data sets in Montana and New Hampshire, USA, that exemplify time-scales at which closure is often assumed. These data sets illustrate different sampling designs that allow testing for closure but are currently rarely employed in field investigations. Using a simulation study, we then evaluate the sensitivity of parameter estimates to changes in site occupancy and evaluate a power analysis developed for sampling designs that is aimed at limiting the likelihood of closure. 3. Application of our approach to point-count data indicates that habitats may frequently be open to changes in site occupancy at time-scales typical of many occupancy investigations, with 71% and 100% of species investigated in Montana and New Hampshire respectively, showing violation of closure across time periods of 3 weeks and 8 days respectively. 4. Simulations suggest that models assuming closure are sensitive to changes in occupancy. Power analyses further suggest that the modelling procedure we apply can effectively test for closure. 5. Synthesis and applications. Our demonstration that sites may be open to changes in site occupancy over time-scales typical of many occupancy investigations, combined with the sensitivity of models to violations of the closure assumption, highlights the importance of properly addressing the closure assumption in both sampling designs and analysis. Furthermore, inappropriately applying closed models could have negative consequences when monitoring rare or declining species for conservation and management decisions, because violations of closure typically lead to overestimates of the probability of occurrence.

To investigate the behavior of the lithosphere undergoing extension, we use a simple rheological model broadly consistent with experimental data on rock creep and with the nature of the brittle/ductile transition. A plastic surface layer overlies a substrate that deforms by power law creep with a stress exponent n = 3 and an effective viscosity that decreases with depth. In extension this model shows a strong necking instability, provided that the thermal gradient is sufficiently large; otherwise, stable uniform extension is indicated. The predicted structures display uniformly spaced necks or regions of enhanced extension (basins) alternating with regions of reduced extension (ranges). If the depth to the brittle/ductile transition is roughly 10 km, as suggested by the maximum depth of seismic faulting, the model yields spacings for the incipient Basin and Range structures of about 25–60 km, in excellent agreement with observation.

Spheroidal weathering, a common mechanism that initiates the transformation of bedrock to saprolite, creates concentric fractures demarcating relatively unaltered corestones and progressively more altered rindlets. In the spheroidally weathering Rio Blanco quartz diorite (Puerto Rico), diffusion of oxygen into corestones initiates oxidation of ferrous minerals and precipitation of ferric oxides. A positive ∆V of reaction results in the buildup of elastic strain energy in the rock. Formation of each fracture is postulated to occur when the strain energy in a layer equals the fracture surface energy. The rate of spheroidal weathering is thus a function of the concentration of reactants, the reaction rate, the rate of transport, and the mechanical properties of the rock. Substitution of reasonable values for the parameters involved in the model produces results consistent with the observed thickness of rindlets in the Rio Icacos bedrock (≈ 2-3 cm) and a time interval between fractures (≈ 200-300 a) based on an assumption of steady-state denudation at the measured rate of 0.01 cm/a. Averaged over times longer than this interval, the rate of advance of the bedrock-saprolite interface during spheroidal weathering (the weathering advance rate) is constant with time. Assuming that the oxygen concentration at the bedrock-saprolite interface varies with the thickness of soil/saprolite yields predictive equations for how weathering advance rate and steadystate saprolite/soil thickness depend upon atmospheric oxygen levels and upon denudation rate. The denudation and weathering advance rates at steady state are therefore related through a condition on the concentration of porewater oxygen at the base of the saprolite. In our model for spheroidal weathering of the Rio Blanco quartz diorite, fractures occur every ~ 250 years, ferric oxide is fully depleted over a four rindlet set in ~ 1000 years, and saprolitization is completed in ~ 5000 years in the zone 2 containing ~ 20 rindlets. Spheroidal weathering thus allows weathering to keep up with the high rate of denudation by enhancing access of bedrock to reactants by fracturing.Coupling of denudation and weathering advance rates can also occur for the case that weathering occurs without spheroidal fractures, but for the same kinetics and transport parameters, the maximum rate of saprolitization achieved would be far smaller than the rate of denudation for the Rio Blanco system. The spheroidal weathering model provides a quantitative picture of how physical and chemical processes can be coupled explicitly during bedrock alteration to soil to explain weathering advance rates that are constant in time.

It has been hypothesized that many soil profi les reach a steady-state thickness. In this work, such profi les were simulated using a one-dimensional model of reaction with advective and diffusive solute transport. A model 'rock' is considered, consisting of albite that weathers to kaolinite in the presence of chemically inert quartz. The model yields three different steadystate regimes of weathering. At the lowest erosion rates, a local-equilibrium regime is established where albite is completely depleted in the weathering zone. This regime is equivalent to the transport-limited regime described in the literature. With an increase in erosion rate, transition and kinetic regimes are established. In the transition regime, both albite and kaolinite are present in the weathering zone, but albite does not persist to the soil-air interface. In the weathering-limited regime, here called the kinetic regime, albite persists to the soil-air interface. The steady-state thickness of regolith decreases with increasing erosion rate in the local equilibrium and transition regimes, but in the kinetic regime, this thickness is independent of erosion rate. Analytical expressions derived from the model are used to show that regolith production rates decrease exponentially with regolith thickness. The steady-state regolith thickness increases with the Darcy velocity of the pore fl uid, and in the local equilibrium regime may vary markedly with small variations in this velocity and erosion rate. In the weathering-limited regime, the temperature dependences for chemical weathering rates are related to the activation energy for the rate constant for mineral reaction and to the ΔH of dissolution, while for local equilibrium regimes they are related to the ΔH only. The model illustrates how geochemical and geomorphological observations are related for a simple compositional system. The insights provided will be useful in interpreting natural regolith profi les.