Choosing appropriate temporal and spatial scales for ecological restoration

Classic ecological restoration seems tacitly to have taken the Clementsian “balance of nature” paradigm for granted: plant succession terminates in a climax community which remains at equilibrium until exogenously disturbed after which the process of succession is restarted until the climax is reached. Human disturbance is regarded as unnatural and to have commenced in the Western Hemisphere at the time of European incursion. Classic ecological restoration thus has a clear and unambiguous target and may be conceived as aiming to foreshorten the natural processes that would eventually lead to the climax of a given site, which may be determined by its state at “settlement”. According to the new “flux of nature” paradigm in ecology a given site has notelos and is constantly changing. Human disturbance is ubiquitous and long-standing, and at certain spatial and temporal scales is “incorporated”. Any moment in the past 10,000 years that may be selected as a benchmark for restoration efforts thus appears to be arbitrary. Two prominent conservationists have therefore suggested that the ecological conditions in North America at the Pleistocene—Holocene boundary, prior to the anthropogenic extinction of the Pleistocene megafauna, be the target for ecological restoration. That suggestion explicitly assumes evolutionary temporal scales and continental spatial scales as the appropriate frame of reference for ecological restoration. However, ecological restoration should be framed in ecological spatio-temporal scales, which may be defined temporally in reference to ecological processes such as disturbance regimes and spatially in reference to ecological units such as landscapes, ecosystems, and biological provinces. Ecological spatio-temporal scales are also useful in achieving a scientifically defensible distinction between native and exotic species, which plays so central a role in the practice of ecological restoration and the conservation of biodiversity. Because post-settlement human disturbances have exceeded the limits of such scales, settlement conditions can be justified scientifically as appropriate targets of restoration efforts without recourse to obsolete teleological concepts of equilibria and without ignoring the presence and ecological influence of indigenous peoples.     

Characteristic spatial and temporal scales unify models of animal movement

Animal movements have been modeled with diffusion at large scales and with more detailed movement models at smaller scales. We argue that the biologically relevant behavior of a wide class of movement models can be efficiently summarized with two parameters: the characteristic temporal and spatial scales of movement. We define these scales so that they describe movement behavior both at short scales (through the velocity autocorrelation function) and at long scales (through the diffusion coefficient). We derive these scales for two types of commonly used movement models: the discrete-step correlated random walk, with either constant or random step intervals, and the continuous-time correlated velocity model. For a given set of characteristic scales, the models produce very similar trajectories and encounter rates between moving searchers and stationary targets. Thus, we argue that characteristic scales provide a unifying currency that can be used to parameterize a wide range of ecological phenomena related to movement.     

Plant reproductive allocation predicts herbivore dynamics across spatial and temporal scales

Life-history theory suggests that iteroparous plants should be flexible in their allocation of resources toward growth and reproduction. Such plasticity could have consequences for herbivores that prefer or specialize on vegetative versus reproductive structures. To test this prediction, we studied the response of the cactus bug (Narnia pallidicornis) to meristem allocation by tree cholla cactus (Opuntia imbricata). We evaluated the explanatory power of demographic models that incorporated variation in cactus relative reproductive effort (RRE; the proportion of meristems allocated toward reproduction). Field data provided strong support for a single model that defined herbivore fecundity as a time-varying, increasing function of host RRE. High-RRE plants were predicted to support larger insect populations, and this effect was strongest late in the season. Independent field data provided strong support for these qualitative predictions and suggested that plant allocation effects extend across temporal and spatial scales. Specifically, late-season insect abundance was positively associated with interannual changes in cactus RRE over 3 years. Spatial variation in insect abundance was correlated with variation in RRE among five cactus populations across New Mexico. We conclude that plant allocation can be a critical component of resource quality for insect herbivores and, thus, an important mechanism underlying variation in herbivore abundance across time and space.     

Interference at several temporal and spatial scales between two chestnut insects

Detection of interspecific competition between insects is often sensitive to scaling. We give an example of scale-dependent interference between the weevil Curculio elephas and the moth Cydia splendana, which both have larvae that develop in the fruits of chestnut Castanea sativa. Measures at three scales were considered: chestnut, husk (with one to three fertile fruits) and tree. Data come from observations in the field over 14 years, complemented by experiments done directly in trees. Data on individual chestnut fruits revealed a marked statistical interference between the two insects. Experiments demonstrated that presence of a moth larva in a fruit usually inhibits weevil egg-laying. Conversely, weevil presence does not strongly modify moth larval behavior. Cases of double infestation often correspond to fruits first attacked by the weevil. With measures on husks, interference between the two insects was observed only in some trees; its intensity was always weaker than in the chestnuts themselves. At the scale of entire trees, rates of infestation by each insect are not correlated. Interference in chestnut fruits is interpreted by assuming that the weevil female either is sensitive to a repellent molecule originating from a moth larva or its frass, or can detect moth larval sounds. Mechanisms governing infestation rates from data per tree are discussed in relation to those found at fruit scale and to plant-insect interactions. The need to estimate available resources both from quantitative and qualitative points of view is emphasized.     

A test of spatial memory and movement patterns of bumblebees at multiple spatial and temporal scales

Naive bumblebee foragers appear to use movement rules at smallspatial and temporal scales, but it is not clear whether theserules determine movement patterns as the scales increase. Onestrategy for efficient foraging used by bumblebees is near-farsearch, involving short flights when in good patches of flowersand longer flights when in poor patches. Bumblebees also demonstratethe use of a spatial memory strategy by returning repeatedlyto patches of flowers, and even following the same route betweenflowers, over periods of days. We attempted to determine atwhat spatial scales bumblebees use spatial memory while foragingwithin a patch and after how many flower visits spatial memoryoutweighs near-far search. Bumblebees in the laboratory foragedon a 4 x 4 array of artificial flowers with distances rangingfrom 10 to 80 cm between flowers in two simple spatial patterns.The proportion of visits to flowers containing a sucrose rewardwas monitored for either 100 or 400 flower visits in two separateexperiments, after which the locations of the rewarding andnonrewarding flowers were interchanged, producing a mirror image.A drop in accuracy after the mirror image switch would indicatethat the bees had memorized the location of rewarding flowers.Mirror image tests, and comparisons to a simulation model ofnear-far search based on actual flight distances, indicate thatnaive bumblebees used near-far search on flowers 10 cm apartbut increasingly used spatial memory as experience and spatialseparation increased. Bumblebees thus have multiple tacticsavailable to forage efficiently in different environments.     

Interactions between spatial and temporal scales in the evolution of dispersal rate

The evolution of dispersal rate is studied with a model of several local populations linked by dispersal. Three dispersal strategies are considered where all, half or none of the offspring disperse. The spatial scale (number of patches) and the temporal scale (probability of local extinction) of the environment are critical in determining the selective advantage of the different dispersal strategies. The results from the simulations suggest that an interaction between group selection and individual selection results in a different outcome in relation to the spatial and temporal scales of the environment. Such an interaction is able to maintain a polymorphism in dispersal strategies. The maintenance of this polymorphism is also scale-dependent. This study suggests a mechanism for the short-term evolution of dispersal, and provides a testable prediction of this hypothesis, namely that loss of dispersal abilities should be more frequent in spatially more continuous environments, or in temporally more stable environments. This revised version was published online in July 2006 with corrections to the Cover Date.     

Population dynamics and potential of fisheries stock enhancement: practical theory for assessment and policy analysis

The population dynamics of fisheries stock enhancement, and its potential for generating benefits over and above those obtainable from optimal exploitation of wild stocks alone are poorly understood and highly controversial. I review pertinent knowledge of fish population biology, and extend the dynamic pool theory of fishing to stock enhancement by unpacking recruitment, incorporating regulation in the recruited stock, and accounting for biological differences between wild and hatchery fish. I then analyse the dynamics of stock enhancement and its potential role in fisheries management, using the candidate stock of North Sea sole as an example and considering economic as well as biological criteria. Enhancement through release of recruits or advanced juveniles is predicted to increase total yield and stock abundance, but reduce abundance of the naturally recruited stock component through compensatory responses or overfishing. Economic feasibility of enhancement is subject to strong constraints, including trade-offs between the costs of fishing and hatchery releases. Costs of hatchery fish strongly influence optimal policy, which may range from no enhancement at high cost to high levels of stocking and fishing effort at low cost. Release of genetically maladapted fish reduces the effectiveness of enhancement, and is most detrimental overall if fitness of hatchery fish is only moderately compromised. As a temporary measure for the rebuilding of depleted stocks, enhancement cannot substitute for effort limitation, and is advantageous as an auxiliary measure only if the population has been reduced to a very low proportion of its unexploited biomass. Quantitative analysis of population dynamics is central to the responsible use of stock enhancement in fisheries management, and the necessary tools are available.     

Modelling the spatial and temporal variability of diatoms in the River Murray

We have developed a simple mechanistic model, which simulates the temporaland spatial variability of concentrations of the diatomAulacoseira granulata in the River Murray, NSWAustralia, over large distances under different flow conditions. The modelincorporates a simple growth function determined by the availabilities ofsilica and light, and by temperature. It also includes losses due tosinking and downstream advection. The model is tested against weeklymeasurements for the period 1981-1990, at three sites separated by 1100 km.The seasonal cycle of Aulacoseira concentrations alongthe river is well represented by the model. During times of high flow, theconcentrations of Aulacoseira remain fairly uniformalong the river as the diatoms have insufficient time to grow before beingadvected downstream.     

Plant traits respond to the competitive neighbourhood at different spatial and temporal scales

Background and Aims Clonal plants can plastically modify their traits in response to competition, but little is known regarding the spatio-temporal scale at which a competitive neighbourhood determines the variability in species traits. This study tests the hypothesis that the local neighbourhood can be expected to influence the processes that are involved in competition tolerance and avoidance, and that this effect depends on organ lifespan.Methods Fragments of the rhizomatous Elytrigia repens (Poaceae) were sampled in 2012 in experimental plant communities that varied in species identity and abundance. These communities had been cultivated since 2009 in mesocosms in a common garden. Fragment performance, shoot and clonal traits were measured, and the effects of past and present local neighbourhoods of five different radius sizes (5–25 cm) were examined. Past and present local neighbourhood compositions were assessed in 2011 and 2012, respectively.Key Results Most of the measured traits of E. repens responded to the local neighbourhood (5–10 cm radius), with an additional effect of the larger neighbourhood (20–25 cm radius) on ramet height, leaf dry matter content, maximal internode length and specific rhizome mass. Contrary to the expectation of the hypothesis, the temporal influence was not due to the organ lifespan. Indeed, five of the eight traits studied responded to both the past and present neighbourhoods. With the exception of specific rhizome mass, all trait responses were explained by the abundance of specific species.Conclusions This study demonstrates that the traits of a single clonal individual can respond to different competitive environments in space and time. The results thus contribute to the understanding of competition mechanisms.     

Interdependence of plankton spatial distribution and plancton biomass temporal oscillations: mathematical simulation

The dynamics of aquatic biological communities in a patchy environment is of great interest in respect to interrelations between phenomena at various spatial and time scales. To study the complex plankton dynamics in relation to variations of such a biologically essential parameter as the fish predation rate, we use a simple reaction-diffusion model of trophic interactions between phytoplankton, zooplankton, and fish. We suggest that plankton is distributed between two habitats one of which is fish-free due to hydrological inhomogeneity, while the other is fish-populated. We show that temporal variations in the fish predation rate do not violate the strong correspondence between the character of spatial distribution of plankton and changes of plankton biomass in time: regular temporal oscillations of plankton biomass correspond to large-scale plankton patches, while chaotic oscillations correspond to small-scale plankton patterns. As in the case of the constant fish predation rate, the chaotic plankton dynamics is characterized by coexistence of the chaotic attractor and limit cycle.     

How spatial and temporal scales influence understanding of Sperm Whale distribution: a review

For much of the past two centuries, sperm whaling has been economically very profitable, and therefore whalers and, more recently, scientists have tried to understand Sperm Whale ( Physeter macrocephalus ) distribution and the factors controlling it. However, due to their deep-living mode of life, their cosmopolitan distribution and our limited knowledge about their main prey species, the task has proved very difficult. This paper reviews studies of Sperm Whale distribution and the factors controlling it. The conclusions of these studies are very equivocal and often appear to contradict one another. It is suggested that this apparent confusion is mainly due to poorly defined spatial and temporal scales, the use of only one scale in most studies and the absence of consideration of the spatial and temporal scales at which relevant oceanographic processes occur. It is concluded that multiscale studies, covering the scales of oceanographic process, are needed to obtain a better understanding of Sperm Whale distribution.     

Current status and potential contributions of fisheries statistics from artisanal fisheries for managing juvenile istiophorid billfishes in Southern Brazil

Environmental Biology of Fishes - Billfishes are considered important fishery resources and the identification of aggregation sites is imperative for proper management. Here we present evidence of...     

Modelling temporal decoupling between biomass and numbers during the transient nitrogen-limited growth of a marine phytoflagellate

A mathematical model of single-nutrient-limited algal growthis presented in which carbon fixation and cell division aredifferent functions of assimilated nutrient. The model successfullydescribes key features of the growth of Isochrysis galbana inammonium-limited batch culture under continuous illuminationand in light/dark cycles. The incorporation of a nutrient processingtime allows the simulation of a time lag between net carbonfixation and cell division, and enables the model to describechanges in the mean carbon content and carbon/nitrogen ratioof the cells. The model can be completely parameterized fromstandard batch culture experiments.     

The influence of bite size on foraging at larger spatial and temporal scales by mammalian herbivores

Organisms respond to their heterogeneous environment in complex ways at many temporal and spatial scales. Here, I examine how the smallest scale process in foraging by mammalian herbivores, taking a bite, influences plants and herbivores over larger scales. First, because cropping bites competes with chewing them, bite size influences short-term intake rate of herbivores within plant patches. On the other hand, herbivores can chew bites while searching for new ones, thus influencing the time spent vigilant and intake rate as animals move among food patches. Therefore, bite size affects how much time herbivores must spend foraging each day. Because acquiring energy is necessary for fitness, herbivores recognize the importance of bite size and select bites, patches and diets based on tradeoffs between harvesting rates, digestion, and sheering forces. In turn, induced structural defenses of plants, such as thorns, allow plants to respond immediately to herbivory by reducing bite size and thus tissue loss. Over evolutionary time, herbivores have adapted mouth morphology that allows them to maximize bite size on their primary forage plant, whereas plants faced with large mammalian herbivores have adapted structures such as divarication that minimize bite size and protect themselves from herbivory. Finally, bite size available among plant communities can drive habitat segregation and migration of larger herbivores across landscapes.     

Modelling below- and above-ground biomass for non-woody and woody plants

BACKGROUND AND AIMS: Intraspecific relationships between below- and above-ground biomass (MB and MA, respectively) have been studied extensively to evaluate environmental effects on growth and development at the level of the individual plant. However, no current theoretical model for this relationship exists for broad interspecific trends. The aims of this paper are to provide a model and to test its predictions using a recently assembled, large database (1406 data entries for 257 species). METHODS: An allometric model was derived to predict the relationship between MB and MA for non-woody and woody plants based on previously developed scaling relationships for leaf, stem and root standing biomass and annual growth rates. The predictions of this model were tested by comparing the numerical values of predicted scaling exponents (the slopes of log-log regression curves) with those observed for the database. KEY RESULTS AND CONCLUSIONS: For non-woody plants and the juveniles of woody species, the model predicts an isometric scaling relationship (i.e. MB proportional, variant MA). For woody plants, a complex scaling function is predicted. But, for a particular set of biologically reasonable conditions, the model predicts MB proportional, variant MA across woody plants. These predictions accord reasonably well with observed statistical trends when non-woody and woody plants are studied separately (n=1061 and 345 data entries, respectively). Although the reliability of regression formulas to estimate MB based on MA measurements increased with increasing plant size, estimates of MB can be as much as two orders of magnitude off, even when using regression formulas with r2 >0.90 and F >53,000.     

Derivation of regional crop sequences as an indicator for potential GMO dispersal on large spatial scales

A methodological approach is presented which aims to visualise the constraints for crop sequence planning in agriculture in a regional, large-scale context. In particular, the relationship between the scope of oilseed rape cultivation and the overall regional cropping structure, the share of particular farm types and the interactions between single crops have been analysed. The identified constraints have been applied to specify current and regionally typical crop sequences as input data for large-scale ex ante assessments, here exemplary for the genome dispersal risk in the case of GM oilseed rape cultivation.The regional and spatio-temporal variation of crop sequences for oilseed rape was analysed and generalised through a combination of analytical, classification and up-scaling techniques. In order to anticipate and assess the dispersal risks of transgenic oilseed rape, the methodology was tuned on crop sequences, which strongly influence the temporal dispersal of genetically modified oilseed rape. The regional cropping patterns for oilseed rape were analysed for the four northernmost German federal states: Schleswig-Holstein, Mecklenburg-Western Pomerania, Lower Saxony and Brandenburg. For typical regional crop clusters, specific crop sequences were derived, taking into account the constraints between crops and the weights for the particular crops as related to farm type. Real land-use data obtained at particular research sites were used to precisely determine the frequency of the single crops, as well as to discover sub-dominant crop combinations, which may have a high impact on dispersal processes. The introduced methodology stresses the following aspects: (i) reflection of the current situation due to links to periodically updated statistical data, (ii) implementation of the relationships and constraints between the different crops through statistical analyses, (iii) transfer of extensive, spatially limited agricultural data and expert knowledge to a large-scale context and (iv) integration of sub-dominant measures that are highly sensitive for particular processes.     

Shifts from competition to facilitation between a foundation tree and a pioneer shrub across spatial and temporal scales in a semiarid woodland

* Theoretical and empirical research has supported the hypothesis that plant-plant interactions change from competition to facilitation with increasing abiotic stress. However, the consistency of such changes has been questioned in arid and semiarid ecosystems. * During a drought in the semiarid south-western USA, we used observations and a field experiment to examine the interactions between juveniles of a foundation tree (Pinyon pine, Pinus edulis) and a common shrub (Apache plume, Fallugia paradoxa) in replicated areas of high and low stress. * The presence of F. paradoxa reduced P. edulis performance at low-stress sites, but had the opposite effect at high-stress sites. However, the intensity of the interactions depended on temporal variation in climate and age of P. edulis. Both above- and below-ground factors contributed to competition, while only above-ground factors contributed to facilitation. * These results support the hypothesis that interactions can change from competition to facilitation as abiotic stress increases in semiarid environments. A shift from competition to facilitation may be important for the recovery of P. edulis and other foundation species that have experienced large-scale mortality during recent droughts.