The development of a digital terrain model for the geomorphological engineering of the ‘rolling’ foredune of Terschelling,the Netherlands

The foredunes form an important element of the line of defence which protects the low-lying parts of the Netherlands from the sea. The foredune of the eastern part of the Wadden island of Terschelling has been managed as a ‘rolling’ foredune to maximize the amount of sand available in times of emergency. Following a decision of the Dutch Government to maintain the coastline of 1990, this foredune will now be stabilized. A plan is made to reshape the morphology of the foredune according to a geomorphological design. A simulation model was developed to produce a Digital Terrain Model with the required geometrical information. The transformation which is on the macro-level scale can be achieved within the envisaged medium-scale planning period of five years only by applying earth-moving machinery, placing fences or planting sand-trapping vegetation.     

Vegetation development in coastal foredunes in relation to methods of establishing marram grass ( Ammophila arenaria )

In coastal foredunes marram grass (Ammophila arenaria) is used to stabilize windblown sand. The development of traditionally plantedAmmophila into a more natural foredune vegetation may take 5–10 yr. For economic reasons, traditional planting may be replaced by alternative techniques such as planting seeds or disk-harrowing rhizome fragments. In this paper, we compare the initial vegetation development of traditionally planted stands with stands established from seeds and from rhizomes. The experiments were conducted on an artificial foredune originating from dredged sea sand. The total experimental area covered more than 100 ha and the vegetation development was studied for 6 yr. The data were analysed bya priori grouping of plant species according to their ecology, as well as by Principal Components Analysis (PCA) and Redundancy Analysis (RA) of the percentage ground cover per plant species. Comparing ecological groups of plants showed that all planting methods delivered equal numbers of plant species that are indicative for coastal dunes. PCA and RA showed that methods based on the use of rhizome material resulted in a higher percentage cover of clonal perennials (Calammophila baltica, Festuca rubra ssp.arenaria, Carex arenaria andCirsium arvense) than the traditionally planted stands and the stands obtained from seeds. The latter two were characterized by the dominance of annuals, bi-annuals and (mostly nonrhizomatous) perennials. Initially, the rates of succession were highest in the stands obtained from rhizomes. However, after 3–6 yr there were no differences between the various stands. During the first four years, the percentage cover by rhizomatous foredune plants developed faster than that of annuals, bi-annuals and perennials. After 6 yr, the latter contributed almost as much to the percentage cover as the clonal species.     

Vegetation development in coastal foredunes in relation to methods of establishing marram grass (Ammophila arenaria)

In coastal foredunes marram grass (Ammophila arenaria) is used to stabilize windblown sand. The development of traditionally plantedAmmophila into a more natural foredune vegetation may take 5 – 10 yr. For economic reasons, traditional planting may be replaced by alternative techniques such as planting seeds or disk-harrowing rhizome fragments. In this paper, we compare the initial vegetation development of traditionally planted stands with stands established from seeds and from rhizomes. The experiments were conducted on an artificial foredune originating from dredged sea sand. The total experimental area covered more than 100 ha and the vegetation development was studied for 6 yr. The data were analysed bya priori grouping of plant species according to their ecology, as well as by Principal Components Analysis (PCA) and Redundancy Analysis (RA) of the percentage ground cover per plant species. Comparing ecological groups of plants showed that all planting methods delivered equal numbers of plant species that are indicative for coastal dunes. PCA and RA showed that methods based on the use of rhizome material resulted in a higher percentage cover of clonal perennials (Calammophila baltica, Festuca rubra ssp.arenaria, Carex arenaria andCirsium arvense) than the traditionally planted stands and the stands obtained from seeds. The latter two were characterized by the dominance of annuals, bi-annuals and (mostly nonrhizomatous) perennials. Initially, the rates of succession were highest in the stands obtained from rhizomes. However, after 3 – 6 yr there were no differences between the various stands. During the first four years, the percentage cover by rhizomatous foredune plants developed faster than that of annuals, bi-annuals and perennials. After 6 yr, the latter contributed almost as much to the percentage cover as the clonal species. Nomenclature: van der Meijden et al. (1990) for plant species. The plant species were classified into ecological groups according to van der Meijden et al. (1991) and Mennema et al. (1980).     

Global climate change: an opportunity for coastal dunes??

The predictions for coastal change under the scenario of global sea-level rise offer impending disaster for the variety of coastal morphologies, their associated habitats, and the accompanying infrastructure. However, the predictions tend to ignore the role of sediment budget in the maintenance of coastal morphology and the dynamics of sediment transfers in the beach-dune sand-sharing system. Accepting that shoreline displacement may be an outcome of sea-level rise and a negative sediment budget, conditions are presented that could lead to a positive or equilibrium sediment budget in the coastal foredune and the retention of the foredune system even as it is being displaced. Accommodation space is a key requirement for the continued functioning of the foredune morphologies during periods of sea-level rise.     

Inferring linear feature use in the presence of GPS measurement error

Global Positioning System (GPS) collars are increasingly used to study animal movement and habitat use. Measurement error is defined as the difference between the observed and true value being measured. In GPS data measurement error is referred to as location error and leads to misclassification of observed locations into habitat types. This is particularily true when studying habitats of small spatial extent with large amounts of edge, such as linear features (e.g. roads and seismic lines). However, no consistent framework exists to address the effect of measurement error on habitat classification of observed locations and resulting biological inference. We developed a mechanistic, empirically-based method for buffering linear features that minimizes the underestimation of animal use introduced by GPS measurement error. To do this we quantified the distribution of measurement error and derived an explicit formula for buffer radius which incorporated the error distribution, the width of the linear feature, and a predefined amount of acceptable type I error in location classification. In our empirical study we found the GPS measurement error of the Lotek GPS_3300 collar followed a bivariate Laplace distribution with parameter ρ = 0.1123. When we applied our method to a simulated landscape, type I error was reduced by 57%. This study highlights the need to address the effect of GPS measurement error in animal location classification, particularily for habitats of small spatial extent.     

Geospatial analysis of vulnerable beach-foredune systems from decadal time series of lidar data

Time series of lidar data, acquired over the past decade along the North American East Coast, provide opportunities to gain new insights into 3D evolution of barrier islands and their beach and dune systems. GIS-based per grid cell statistics and map algebra was applied to time series of Digital Surface Models representing two sections of North Carolina barrier islands to quantify elevation change trends, map dynamic and stable locations, identify new and lost buildings, measure relative volume evolution in the beach and foredune systems and analyze shoreline dynamics. Results show a relatively small stable core in both study areas, with beaches and the ocean side of the dunes exhibiting systematic high rates of elevation loss while areas landward from the dunes increase slightly in elevation. Significant number of new homes have been built at locations with very small core surface elevation, and homes built within the shoreline dynamics band have already been lost. The raster-based methodology used in this study can be applied to perform similar analyses in other coastal areas where time series of lidar data are available.     

Contrasting approved uses against actual uses at La Restinga Lagoon National Park, Margarita Island, Venezuela. A GPS and GIS method to improve management plans and rangers coverage

Protected areas are the core of efforts to conserve biological diversity and zoning uses, and they are used as a tool for their management. Global Positioning Systems (GPS) and Geographic Information System (GIS) have been used to evaluate conflicts between approved uses and actual uses in La Restinga Lagoon National Park in Venezuela (LRLNP). The park (188.6 km²), covers various ecosystems such as coastal lagoons, marine waters and xerophytic vegetation, it is visited by up to 260,000 people visit per year. A GIS, using a base map compiled from 1:25,000 maps was developed. Natural cover was mapped from LANDSAT VII TM images, orthophotomaps and aerial photographs. Spatial use data was collected by field GPS location of any use inside the park during 2 years. Thematic vector maps for each land cover, zoning area, and observed use were created. Maps of zoning and uses were overlapped and new maps for each use-zoning crossing were created. The park contain nine different zoning areas wherein 28 different uses were identified, 18 of them were direct consumptive and non-consumptive uses, occurring on 54% of the park. Tourism transit areas were the most used zones. More than 5,000 people use the park during a high season day. Superimposition of zoning maps with actual use data produced 13 uses taking place in not allowed areas (46% of park area). Most common prohibited use was commercial net fishing, occurring in nearly 40% of the park area. Therefore, identifying human use conflicts and its geographical distribution is a key issue to improve Management Plans as well as identifying hot sites. The GPS-GIS methodology presented here allows Park Rangers to select those areas which could benefit from enhanced supervision with the limited budget available.     

中国北方农牧交错带农村土地利用变化因子分析--以内蒙古尧勒甸子村为例

以北方农牧交错带典型生态示范村尧勒甸子村为例,应用参与式农村评估法(PRA)并结合GPS与野外测量,恢复了该村1949年后50 a来的土地利用历史过程与格局,利用GIS分析其1950-1999年的土地利用变化特点.运用因子分析和逐步回归分析方法揭示了人类活动因素对土地利用变化的作用.结果发现1985年以前耕地面积扩大,草地面积减小,沙地面积迅猛持续增加;1985年以后土地利用变化向相反的方向发展,土地利用结构趋向合理.     

Spatial and temporal patterns of habitat use by Eurasian oystercatchers (<Emphasis Type="Italic">Haematopus ostralegus</Emphasis>) in the eastern Wadden Sea revealed using GPS data loggers

Detailed knowledge on species–habitat relationships is of crucial importance for the understanding of processes in marine ecosystems. Being top-predators, birds are important bio-indicators for marine systems. The aim of this study was to elucidate precise information on foraging habitat use and foraging times of oystercatchers (Haematopus ostralegus) on wide tidal flats using global positioning system (GPS) data loggers. The study was conducted to collect hints for the negative population trends in oystercatchers in the Wadden Sea. It is the first time that GPS technique has been used in a shorebird species. Although oystercatchers are known to exhibit foraging site fidelity, a number of individuals visited multiple sites. Foraging trips at night were longer, and the targeted sites were further away than those used during the day. These patterns were likely to be caused by higher risks of clutch predation by avian predators during the day that led adults to reduce their absence to defend their clutches. Our methodological approach enabled the subtle spatio-temporal patterns of habitat use to be determined on a very fine spatio-temporal scale. We suggest further potential studies using GPS data loggers that may help to reveal the reasons for the current declines in oystercatcher populations in the German Wadden Sea.     

Information content is more important than sensory system or physical distance in guiding the long-term evolutionary relationships between signaling modalities in Sceloporus lizards

Long-term signal evolution is shaped by a variety of selective pressures including the need to convey additional information or to improve message transfer to specific receivers or through multiple environments. Here, we test the relative importance of information and sensory modality in shaping the long-term evolution of multimodal signals in Sceloporus lizards. To broadcast identity at territorial boundaries, male Sceloporus use both visual motion (headbob) and chemical signals, whereas they use color (blue belly patches) to signal aggression. Using modern phylogenetic comparative methods, we found a negative correlation between evolutionary changes in visual motion (headbobs) and chemical (femoral pore) signals, but only indirect ties between the evolution of color and motion signals (both of which are perceived visually) through viviparity, and no evidence of an evolutionary link between color and chemical signals. We also find a negative correlation between arboreality and chemical signals. Thus, information content (in this case, broadcasting individual identity versus signaling aggression) appears to play a more important role than sensory modality or physical distance in guiding long-term signal evolution. Additional insights into the underlying evolutionary processes are described, illustrating the utility of a phylogenetic approach.     

Social monitoring in a multilevel society: a playback study with male Guinea baboons

Keeping track of social interactions among conspecifics is a driving force for the evolution of social cognition. How social cognition, such as social knowledge, ties in with a species' social organization is, however, largely unexplored. We investigated the social knowledge of wild Guinea baboons (Papio papio) ranging in Senegal, a species that lives in a fluid multilevel society with overlapping habitat use. Using playback experiments, we tested how adult males differentiate between subjects from their own vs. a neighboring or a stranger social unit (“gang”) and assessed ranging patterns with Global Positioning System (GPS) data. While territorial species usually differentiate between group and nongroup members and often respond more strongly to strangers than neighbors (the “dear enemy” effect), subjects in this highly tolerant species should largely ignore other unit members and mainly attend to subjects from their own unit. Males responded strongly after playback of calls recorded from members of their own gang, while they attended only briefly to neighbor or stranger calls. Apparently, males benefit from monitoring the social maneuvers in their own social unit, while it remains to be resolved whether they are unmotivated or unable to keep track of the identities and actions of individuals outside their own gang. The study highlights how the allocation of social attention is tuned to the specifics of a species' social organization, while a complex social organization does not necessarily translate into the need for more elaborate social knowledge.     

GPS tracking a marine predator: the effects of precision,resolution and sampling rate on foraging tracks of African Penguins

We used a prototype GPS logger to track the movements of breeding African Penguins (Spheniscus demersus). The loggers also recorded temperature and water depth, which allowed us to reconstruct foraging tracks in three dimensions, although GPS signals are interrupted when the birds dive. Here we report the loggers performance in the field and assess the effects of GPS error, resolution and sampling rate on estimates of foraging track length and speed. There is a trade-off between sampling rate and battery lifespan. We tested loggers at sampling intervals of 1 s, 10 s, 1 min, 2 min and 10 min. Sampling less frequently increases the chance of tracking an entire foraging trip, but it slows uplink times, slightly decreases the accuracy of positional fixes, and significantly reduces the ability to measure fine-scale aspects of foraging behaviour. Compared with radio or satellite tracking, GPS loggers offer unprecedented detail about animal movements. The results of our analysis suggest that techniques that sample relatively infrequently, such as satellite tracking, underestimate actual track lengths by up to 50%. However, caution is needed when interpreting fine-scale sampling for relatively slow-moving organisms. Re-sampling 1-s tracks suggests that c. 35% of apparent movements at this scale are due to measurement error and, more importantly, the limited spatial resolution of GPS (1.85×1.54 m at the study area). We recommend that researchers use a 1-s sampling rate for fine-scale studies, but resample at less frequent intervals to remove spurious noise for slow-moving animals. At current levels of resolution, animals should move at least 4 m per sampling interval. We provide empirical correction factors to compare inferred track length sampled at different rates, but caution that these are idiosyncratic and strongly dependent on the animals behaviour. Overall, GPS loggers offer a significant advance for studies of fine-scale animal movement patterns.Communicated by O. Kinne, Oldendorf/Luhe     

Behavioral transitions with the evolution of cooperative nest founding by harvester ant queens

Research on the evolution of cooperative groups tends to explore the costs and benefits of cooperation, with less focus on the proximate behavioral changes necessary for the transition from solitary to cooperative living. However, understanding what proximate changes must occur, as well as those pre-conditions already in place, is critical to understanding the origins and evolution of sociality. The California harvester ant Pogonomyrmex californicus demonstrates population-level variation in colony founding over a close geographic range. In adjacent populations, queens either found nests as single individuals (haplometrosis) or form cooperative groups of nonrelatives (pleometrosis). We compared aggregation, aggression, and tolerance of queens from one pleometrotic and two haplometrotic populations during nest initiation, to determine which behaviors show an evolutionary shift and which are present at the transition to pleometrosis. Surprisingly, within-nest aggregative behavior was equally present among all populations. In nesting boxes with multiple available brood-rearing sites, both queen types readily formed and clustered around a single common brood pile, suggesting that innate attraction to brood (offspring) facilitates the transition to social aggregation. In contrast, queens from the three populations differed in their probabilities of attraction on the ground to nest sites occupied by other queens and in levels of aggression. Our results suggest that some key behavioral mechanisms facilitating cooperation in P. californicus are in place prior to the evolution of pleometrosis and that the switch from aggression to tolerance is critical for the evolution of stable cooperative associations.     

When good neighbors don't need fences: temporal landscape partitioning among baboon social groups

Intraspecific competition is a key factor shaping space-use strategies and movement decisions in many species, yet how and when neighbors utilize shared areas while exhibiting active avoidance of one another is largely unknown. Here, we investigated temporal landscape partitioning in a population of wild baboons (Papio cynocephalus). We used global positioning system (GPS) collars to synchronously record the hourly locations of five baboon social groups for ～900 days, and we used behavioral, demographic, and life history data to measure factors affecting use of overlap areas. Annual home ranges of neighboring groups overlapped substantially, as predicted (baboons are considered non-territorial), but home ranges overlapped less when space use was assessed over shorter time scales. Moreover, neighboring groups were in close spatial proximity to one another on fewer days than predicted by a null model, suggesting an avoidance-based spacing pattern. At all time scales examined (monthly, biweekly, and weekly), time spent in overlap areas was greater during time periods when groups fed on evenly dispersed, low-quality foods. The percent of fertile females in social groups was negatively correlated with time spent in overlap areas only during weekly time intervals. This suggests that broad temporal changes in ecological resources are a major predictor of how intensively overlap areas are used, and groups modify these ecologically driven spacing patterns at short time scales based on female reproductive status. Together, these findings offer insight into the economics of territoriality by highlighting the dynamics of spacing patterns at differing time scales.     

Size and spatial arrangement of home range of checkered snapper <Emphasis Type="Italic">Lutjanus decussatus</Emphasis> (Lutjanidae) in an Okinawan coral reef determined using a portable GPS receiver

Although snappers (Lutjanidae) are commercially important fishery resources in tropical and subtropical waters, their home range size and its spatial arrangement have not been sufficiently clarified. In the present study, the size and spatial arrangement of the home range of the reef-associated checkered snapper Lutjanus decussatus (Lutjanidae) were investigated by use of a portable GPS receiver. In a 120 m × 120 m quadrat established on a section of the fringing reef of Ishigaki Island, 29 individuals of the species were identified by their color pattern and individuals were divided into five arbitrary size classes (class 1 <10 cm TL ≤ class 2 <15 cm TL ≤ class 3 <20 cm TL ≤ class 4 <25 cm TL ≤ class 5). Fish tracking by snorkeling was conducted with the portable GPS receiver. Site fidelity of this species was high. Home range size ranged from 93.0 to 3638.4 m², and there was a significant positive correlation between the home range size and fish total length. Home ranges of the same-sized individuals abutted each other (8.8% area overlap), whereas those of different-sized individuals overlapped (44.0% area overlap). Agonistic behavior (attack and agonistic display) was more frequently found among same-sized individuals (times of agonistic behavior/times of all encounters × 100 = 71.3%), whereas such agonistic behavior was rarely found among different-sized individuals (times of agonistic behavior/times of all encounters × 100 = 6.9%). These results suggest that home ranges of Lutjanus decussatus can be regarded as territories against same-sized individuals, but not different-sized individuals. The usefulness of the fish tracking by snorkeling using a portable GPS receiver for home range size estimation and the function of the overlapping territory of the species are discussed.     

Evolution of spit morphology: a case study using a remote sensing and statistical based approach

The dynamics and factors responsible for morphological changes of spits viz., Uliyargoli-Padukere, Oddu Bengre and Kodi Bengre, southern Karnataka, India, are investigated using multi-dated satellite images and topographic maps during the last 95-years (1910–2005). Variations and overall rate of changes in length, area and coefficient of determination (R²) of each spit are calculated separately for two periods (1910 and 1967 as base years) to find out whether there is any significant trend in the case of change in length and area in all the three spits that are under study. Linear trend lines are fitted using a least squares method and the statistical significance is considered at 80 % level of confidence. The results recorded significant changes in spit morphology, especially in length and area if 1910 and 1967 are considered separately as base years, are may be due to non-availability of data set between 1910 and 1967 period. The study reveals that coastal processes, such as SW-monsoon influenced strong currents and longshore drifts are the main process for formation and growth of spits, whereas rivers influence/drift also plays significant role. The statistical uncertainty estimation in spits morphology is prevalent wherever the coast is affected by human interventions. The study demonstrates that combined use of satellite imagery and statistical techniques can be effectively used to understand the evolution of spits morphology.     

Information processing in models for density-dependent emigration: A comparison

Density-dependent emigration has been recognized as a fitness enhancing strategy. Yet, especially in the modelling literature there is no consensus about how density-dependent emigration should quantitatively be incorporated into metapopulation models. In this paper we compare the performance of five different dispersal strategies (defined by the functional link between density and emigration probability). Four of these strategies are based on published functional relationships between local population density and emigration probability, one assumes density-independent dispersal. We use individual-based simulations of time-discrete metapopulation dynamics and conduct evolution experiments for a broad range of values for dispersal mortality and environmental stochasticity. For each set of these conditions we analyze the evolution of emigration rates in ‘monoculture experiments’ (with only one type of dispersal strategy used by all individuals in the metapopulation) as well as in selection experiments that allow a pair-wise comparison of the performance of each functional type. We find that a single-parameter ‘asymptotic threshold’ strategy - derived from the marginal value theorem - with a decelerating increase of emigration rate with increasing population density, out-competes any other strategy, i.e. density-independent emigration, a ‘linear threshold’ strategy and a flexible three-parameter strategy. Only when environmental conditions select for extremely high emigration probabilities (close to one), strategies may perform approximately equally. A simple threshold strategy derived for the case of continuous population growth performs even worse than the density-independent strategy. As the functional type of the dispersal function implemented in metapopulation models may severely affect predictions concerning the survival of populations, range expansion, or community changes we clearly recommend to carefully select adequate functions to model density-dependent dispersal.     

Modeling a spatially restricted distribution in the Neotropics: How the size of calibration area affects the performance of five presence-only methods

We here examine species distribution models for a Neotropical anuran restricted to ombrophilous areas in the Brazilian Atlantic Forest hotspot. We extend the known occurrence for the treefrog Hypsiboas bischoffi (Anura: Hylidae) through GPS field surveys and use five modeling methods (BIOCLIM, DOMAIN, OM-GARP, SVM, and MAXENT) and selected bioclimatic and topographic variables to model the species distribution. Models were first trained using two calibration areas: the Brazilian Atlantic Forest (BAF) and the whole of South America (SA). All modeling methods showed good levels of predictive power and accuracy with mean AUC ranging from 0.77 (BIOCLIM/BAF) to 0.99 (MAXENT/SA). MAXENT and SVM were the most accurate presence-only methods among those tested here. All but the SVM models calibrated with SA predicted larger distribution areas when compared to models calibrated in BAF. OM-GARP dramatically overpredicted the species distribution for the model calibrated in SA, with a predicted area around 10⁶ km² larger than predicted by other SDMs. With increased calibration area (and environmental space), OM-GARP predictions followed changes in the environmental space associated with the increased calibration area, while MAXENT models were more consistent across calibration areas. MAXENT was the only method that retrieved consistent predictions across calibration areas, while allowing for some overprediction, a result that may be relevant for modeling the distribution of other spatially restricted organisms.     

A mechanistic model of GPS collar location data: Implications for analysis and bias mitigation

Global positioning system (GPS) collars have revolutionized the collection of animal location data; however, it is well-recognized that considerable bias can be present in these data due to habitat or behavior-induced obstruction of satellite signals resulting in inaccurate or missing locations. To date, no explicit theoretical framework of GPS fix acquisition specific to animal telemetry has been presented, and studies make differing assumptions regarding factors influencing GPS fix acquisition and how these data should be analyzed. Inappropriate statistical models have been used, interaction effects have been misunderstood, and the implementation of bias mitigation techniques has been problematic. Herein we outline current conceptual and analytical problems in the GPS animal telemetry literature, and subsequently present a theoretical model-based framework for GPS fix acquisition that clarifies the single and interactive effects of habitat and behavioral obstruction, fix interval, and collar model on GPS collar performance. By recognizing that GPS fix acquisition is a Bernoulli process, it becomes apparent that all forms of obstruction inherently interact with each other, making generalizations across study areas, study species, and collar models problematic. Stationary collar tests to determine the probability of fix acquisition (PFA), location accuracy, and the response to sources of obstruction are thus of limited applicability to animal-deployed collars. Bias mitigation techniques that extrapolate PFA models across samples, especially those using stationary collar tests to correct animal-deployed collars, are theoretically unsound. It is also demonstrated that nonlinearities in the relationships between sources of obstruction and PFA complicate PFA modeling with limited data and that even slight model misspecification can lead to considerable errors in correction factors, especially when using inverse weighting to mitigate bias. By emphasizing the importance of GPS collar sensitivity and ephemeris retention, the theoretical framework predicts that newer, more sensitive GPS collars will be less severely biased by sources of obstruction than reported for the older, less sensitive collars that have been used in the majority of GPS performance studies to date and we expect this trend to continue. This heuristic modeling exercise should be of value to researchers planning and analyzing studies using GPS collars and it also establishes a starting point for future theoretical investigations into GPS collar performance and bias mitigation.     

Elk decision-making rules are simplified in the presence of wolves

The risk of predation drives many behavioral responses in prey. However, few studies have directly tested whether predation risk alters the way other variables influence prey behavior. Here we use information theory (Akaike’s Information Criterion, AICc) in a novel way to test the hypothesis that the decision-making rules governing elk behavior are simplified by the presence of wolves. With elk habitat use as the dependent variable, we test whether the number of independent variables (i.e., the size of the models) that best predict this behavior differ when wolves are present vs absent. Thus, we use AICc scores simply to determine the number of variables to which elk respond when making decisions. We measured habitat use using 2,288 locations from GPS collars on 14 elk, over two winters (14 elk winters), in the Gallatin Canyon portion of the Greater Yellowstone Ecosystem. We found that the use of three major habitat components (grass, conifer, sage) was sensitive to many variables on days that wolves were locally absent, with the best models (ΔAICc≤2) averaging 7.4 parameters. In contrast, habitat use was sensitive to few variables on days when wolves were present: the best models averaged only 2.5 parameters. Because fewer variables affect elk behavior in the presence of wolves, we conclude that elk use simpler decision-making rules in the presence of wolves. This simplification of decision-making rules implies that predation risk imposes selection pressures that do not allow prey to respond to other pressures in ways that they otherwise would. If the affected processes are important, then this indirect effect of predation is likely to be important.