Local resource availability drives habitat use by a threatened avian granivore in savanna woodlands.

Conserving threatened species relies on an understanding of their habitat requirements. This is especially relevant for granivorous birds, whose habitat use and movement patterns are intricately linked to the spatial and temporal availability of resources such as food and water. In this study, we investigated the habitat use, home range and daily activity patterns of the Endangered Southern Black-throated Finch (SBTF; Poephila cincta cincta) within a 75,000 ha savanna woodland study area in northeastern Australia. This semi-arid region is one of the key remaining strongholds for the species and is characterised by substantially different habitat and climatic conditions than areas where previous research on this species has been undertaken. We radio tracked 142 SBTF using both manual tracking and an array of 27 automated radio towers, which revealed a strong preference for eucalypt-dominated grassy woodland communities. The preference for these habitats also increased with decreasing distance to permanent water. While SBTF occupied large home ranges, individual SBTF were largely sedentary during the radio tracking period (21.8, range = 0.83-120 days), with few landscape-scale movements of more than 4.5 km. Daily foraging activity of SBTF exhibited bimodal peaks in the early morning and late afternoon, while other activities were greatest from the late morning to the early afternoon. Compared to other estrildid finches, our research suggests that SBTF track resources at a local scale across a large home range. We postulate that in times of resource scarcity SBTF may use dietary diversification, instead of landscape or regional-scale nomadic movements, to meet their resource needs. The species' movement patterns underscore the importance of local scale habitat management to facilitate resource availability throughout the year. Furthermore, our research helps target monitoring designs for granivorous birds that focus on the species' diurnal activity patterns.

Marine Fish Movement: home range sizes for commercially relevant species.

Estimates of home range sizes for marine fishes are essential for designing and assessing the effects of spatial wildlife conservation policies and management interventions. However, in situ studies of marine species movement are challenging and often expensive, resulting in a paucity of data on the home range size of the vast majority of marine fishes. Here, we develop a set of new datasets, which we have collectively named Marine Fish Movement, that synthesises published empirically evaluated home ranges reported for adult marine fishes that interact with fisheries and leverage these data to estimate home range sizes for unstudied species. The empirical data contain estimated home range sizes (km2) for 193 species across 63 family groups from 179 studies published between 1971 and 2022. We use a random forest regression model to estimate home range sizes (km2) for 664 fished marine species currently lacking home range estimates. Marine Fish Movement can inform spatial interventions including the design and management of marine protected areas and dynamic fisheries management to meet sustainability goals.

The social and defensive function of olfactory behaviors in a pair-living sexually monogamous primate.

Olfactory behaviors serve a wide variety of social functions in mammals. Odor may signal information about attributes of individuals important for mating and reproduction. Olfactory behaviors, such as scent-marking, may also function as part of home range or resource defense strategies. We assessed the potential social and home range defense functions of olfactory behavior in a pair-living and sexually monogamous primate, Azara's owl monkey (Aotus azarae), in the Argentinian Chaco. This is the most extensive investigation of owl monkey olfactory behaviors in the wild. Individuals regularly performed olfactory behaviors (group mean + SD = 1.3 + 0.5 per hour). The patterns were generally comparable to those observed in studies of captive owl monkeys, except that urine washing was the most common behavior in the wild, as opposed to scent-marking and genital inspections. Most olfactory behaviors were performed by adults, and there were striking sex differences in genital inspections: almost all consisted of an adult male inspecting the paired adult female. These findings suggest that olfactory behaviors play an important role in signaling and coordinating reproduction among owl monkeys, particularly during periods of female conception and pregnancy. Additionally, our research indicates that these behaviors may also serve as a defense strategy for maintaining the core area of their home ranges. This study offers the first assessment of the role of olfactory behaviors in reproductive contexts and home range defense in pair-living, monogamous platyrrhine primates.

Use of acoustic signals in Cape fur seal mother-pup reunions: individual signature, signal propagation and pup home range.

The Cape fur seal (Arctocephalus pusillus pusillus) is one of the most colonial mammals, with colonies of up to hundreds of thousands of individuals during the breeding season. During the lactation period, mothers and pups are regularly separated as females undertake multi-day foraging trips at sea. Mothers and pups use a mutual vocal recognition system to reunite after separation. Such communication is highly constrained by both high background noise and risk of individual confusion owing to the density of seals. This study aimed to experimentally assess the acoustic features relevant for mother-pup vocal identification and the propagation properties of their calls. Playback experiments revealed that mother and pup individual vocal signatures rely on both temporal and frequency parameters: amplitude and frequency modulations, timbre and fundamental frequency (f0). This is more parameters than in any colonial species studied so far. The combinational use of acoustic features reinforces the concept that both environmental and social constraints may have acted as selective pressures on the individual vocal recognition systems. Theoretical propagation distances of mother and pup vocalisations were estimated to be below the range of distances at which mother-pup reunions can occur. This suggests that Cape fur seals may have strong abilities to extract vocal signals from the background noise, as previously demonstrated in the highly colonial king penguin. Investigating the transmission of information throughout the propagation of the signal as well as the ability of the receiving individual to decipher vocal signatures is crucial to understanding vocal recognition systems in the wild.

No evidence for the consistent effect of supplementary feeding on home range size in terrestrial mammals.

Food availability and distribution are key drivers of animal space use. Supplemental food provided by humans can be more abundant and predictable than natural resources. It is thus believed that supplementary feeding modifies the spatial behaviour of wildlife. Yet, such effects have not been tested quantitatively across species. Here, we analysed changes in home range size owing to supplementary feeding in 23 species of terrestrial mammals using a meta-analysis of 28 studies. Additionally, we investigated the moderating effect of factors related to (i) species biology (sex, body mass and taxonomic group), (ii) feeding regimen (duration, amount and purpose), and (iii) methods of data collection and analysis (source of data, estimator and spatial confinement). We found no consistent effect of supplementary feeding on changes in home range size. While an overall tendency of reduced home range was observed, moderators varied in the direction and strength of the trends. Our results suggest that multiple drivers and complex mechanisms of home range behaviour can make it insensitive to manipulation with supplementary feeding. The small number of available studies stands in contrast with the ubiquity and magnitude of supplementary feeding worldwide, highlighting a knowledge gap in our understanding of the effects of supplementary feeding on ranging behaviour.

Resource manipulation reveals interactive phenotype-dependent foraging in free-ranging lizards.

Recent evidence suggests that individuals differ in foraging tactics and this variation is often linked to an individual's behavioural type (BT). Yet, while foraging typically comprises a series of search and handling steps, empirical investigations have rarely considered BT-dependent effects across multiple stages of the foraging process, particularly in natural settings. In our long-term sleepy lizard (Tiliqua rugosa) study system, individuals exhibit behavioural consistency in boldness (measured as an individual's willingness to approach a novel food item in the presence of a threat) and aggressiveness (measured as an individual's response to an 'attack' by a conspecific dummy). These BTs are only weakly correlated and have previously been shown to have interactive effects on lizard space use and movement, suggesting that they could also affect lizard foraging performance, particularly in their search behaviour for food. To investigate how lizards' BTs affect their foraging process in the wild, we supplemented food in 123 patches across a 120-ha study site with three food abundance treatments (high, low and no-food controls). Patches were replenished twice a week over the species' entire spring activity season and feeding behaviours were quantified with camera traps at these patches. We tracked lizards using GPS to determine their home range (HR) size and repeatedly assayed their aggressiveness and boldness in designated assays. We hypothesised that bolder lizards would be more efficient foragers while aggressive ones would be less attentive to the quality of foraging patches. We found an interactive BT effect on overall foraging performance. Individuals that were both bold and aggressive ate the highest number of food items from the foraging array. Further dissection of the foraging process showed that aggressive lizards in general ate the fewest food items in part because they visited foraging patches less regularly, and because they discriminated less between high and low-quality patches when revisiting them. Bolder lizards, in contrast, ate more tomatoes because they visited foraging patches more regularly, and ate a higher proportion of the available tomatoes at patches during visits. Our study demonstrates that BTs can interact to affect different search and handling components of the foraging process, leading to within-population variation in foraging success. Given that individual differences in foraging and movement will influence social and ecological interactions, our results highlight the potential role of BT's in shaping individual fitness strategies and population dynamics.

Individual energetics scale up to community coexistence: Movement, metabolism and biodiversity dynamics in fragmented landscapes.

Unravelling the intricate mechanisms that govern community coexistence remains a daunting challenge, particularly amidst ongoing environmental change. Individual physiology and metabolism are often studied to understand the response of individual animals to environmental change. However, this perspective is currently largely lacking in community ecology. We argue that the integration of individual metabolism into community theory can offer new insights into coexistence. We present the first individual-based metabolic community model for a terrestrial mammal community to simulate energy dynamics and home range behaviour in different environments. Using this model, we investigate how ecologically similar species coexist and maintain their energy balance under food competition. Only if individuals of different species are able to balance their incoming and outgoing energy over the long-term will they be able to coexist. After thoroughly testing and validating the model against real-world patterns such as of home range dynamics and field metabolic rates, we applied it as a case study to scenarios of habitat fragmentation - a widely discussed topic in biodiversity research. First, comparing single-species simulations with community simulations, we find that the effect of habitat fragmentation on populations is strongly context-dependent. While populations of species living alone in the landscape were mostly positively affected by fragmentation, the diversity of a community of species was highest under medium fragmentation scenarios. Under medium fragmentation, energy balance and reproductive investment were also most similar among species. We therefore suggest that similarity in energy balance among species promotes coexistence. We argue that energetics should be part of community ecology theory, as the relative energetic status and reproductive investment can reveal why and under what environmental conditions coexistence is likely to occur. As a result, landscapes can potentially be protected and designed to maximize coexistence. The metabolic community model presented here can be a promising tool to investigate other scenarios of environmental change or other species communities to further disentangle global change effects and preserve biodiversity.

Determinants of macaques' space use: A test for the ecological constraints model using GPS collars.

As a central topic in Behavioral Ecology, animal space use involves dynamic responses to social and ecological factors. We collared 22 rhesus macaques (Macaca mulatta) from six groups on Neilingding Island, China, and collected 80,625 hourly fixes over a year. Using this high-resolution location data set, we quantified the macaques' space use at the individual level and tested the ecological constraints model while considering various environmental and human interfering factors. As predicted by the ecological constraints model, macaques in larger groups had longer daily path lengths (DPLs) and larger home ranges. We found an inverted U-shape relationship between mean daily temperatures and DPLs, indicating that macaques traveled farther on mild temperature days, while they decreased DPLs when temperatures were too high or too low. Anthropogenic food subsidies were positively correlated to DPLs, while the effect of rainfall was negative. Macaques decreased their DPLs and core areas when more flowers and less leaves were available, suggesting that macaques shifted their space use patterns to adapt to the seasonal differences in food resources. By applying GPS collars on a large number of individuals living on a small island, we gained valuable insights into within-group exploitation competition in wild rhesus macaques.

Winter GPS tagging reveals home ranges during the breeding season for a boreal-nesting migrant songbird, the Golden-crowned Sparrow.

Determining space use for species is fundamental to understanding their ecology, and tracking animals can reveal insights into their spatial ecology on home ranges and territories. Recent technological advances have led to GPS-tracking devices light enough for birds as small as ~30 g, creating novel opportunities to remotely monitor fine-scale movements and space use for these smaller species. We tested whether miniaturized GPS tags can allow us to understand space use of migratory birds away from their capture sites and sought to understand both pre-breeding space use as well as territory and habitat use on the breeding grounds. We used GPS tags to characterize home ranges on the breeding grounds for a migratory songbird with limited available breeding information, the Golden-crowned Sparrow (Zonotrichia atricapilla). Using GPS points from 23 individuals across 26 tags (three birds tagged twice), we found home ranges in Alaska and British Columbia were on average 44.1 ha (95% kernel density estimate). In addition, estimates of territory sizes based on field observations (mean 2.1 ha, 95% minimum convex polygon [MCP]) were three times smaller than 95% MCPs created using GPS tags (mean 6.5 ha). Home ranges included a variety of land cover classes, with shrubland particularly dominant (64-100% of home range cover for all but one bird). Three birds tracked twice returned to the same breeding area each year, supporting high breeding site fidelity for this species. We found reverse spring migration for five birds that flew up to 154 km past breeding destinations before returning. GPS-tracking technology allowed for critical ecological insights into this migratory species that breeds in very remote locations.

Home range, habitat use and capture-release of translocated leopards in Gir landscape, Gujarat, India.

Understanding the spatial ecology of translocated leopards (Panthera pardus fusca) is crucial for their conservation and the effective assessment of conflict management strategies. We investigated the home range and habitat preferences of five radio-collared leopards (n = 5; 2 males; 3 females) in the Gir landscape. Additionally, we examined the usefulness of the capture-release strategy for these animals. We assessed home range and habitat selection using kernel density estimation (at 95% and 50% levels) and compositional analysis. Our findings revealed that leopards exhibited distinct patterns of movement, often returning to their original capture site or nearby locations or exploring new areas within 3 to 25 days, covering distances ranging from 48 to 260 km. The average home range (95% FK) was estimated at 103.96±36.37 (SE) km2, with a core area usage (50% FK) of 21.38±5.95 km2. Seasonally, we observed the largest home ranges during summer and the smallest during winter. Males exhibited larger home ranges (95% FK, 151±64.28 km2) compared to females (56.18±14.22 km2). The habitat analysis indicated that agricultural areas were consistently preferred in the multi-use landscape at the 2nd order habitat selection level. Additionally, habitat around water bodies was highly favoured at the 3rd order, with distinct variations in habitat selection observed during day and night. This study highlights the significance of riverine and scrubland habitats, as leopards exhibited strong preferences for these habitats within their home ranges. We emphasize the importance of conserving natural habitat patches, particularly those surrounding water bodies. We also report on the characteristics of the capture-release strategy and provide our observations indicating no escalated aggression by leopards' post-release. In conclusion, this study evaluates widely employed approaches to conflict mitigation and suggests the continuous review and assessment of management strategies for mitigating human-leopard conflicts.

Cumulative route improvements spontaneously emerge in artificial navigators even in the absence of sophisticated communication or thought.

Homing pigeons (Columba livia) navigate by solar and magnetic compass, and fly home in idiosyncratic but stable routes when repeatedly released from the same location. However, when experienced pigeons fly alongside naive counterparts, their path is altered. Over several generations of turnover (pairs in which the most experienced individual is replaced with a naive one), pigeons show cumulative improvements in efficiency. Here, I show that such cumulative route improvements can occur in a much simpler system by using agent-based simulation. Artificial agents are in silico entities that navigate with a minimal cognitive architecture of goal-direction (they know roughly where the goal is), social proximity (they seek proximity to others and align headings), route memory (they recall landmarks with increasing precision), and continuity (they avoid erratic turns). Agents' behaviour qualitatively matched that of pigeons, and quantitatively fitted to pigeon data. My results indicate that naive agents benefitted from being paired with experienced agents by following their previously established route. Importantly, experienced agents also benefitted from being paired with naive agents due to regression to the goal: naive agents were more likely to err towards the goal from the perspective of experienced agents' memorised paths. This subtly biased pairs in the goal direction, resulting in intergenerational improvements of route efficiency. No cumulative improvements were evident in control studies in which agents' goal-direction, social proximity, or memory were lesioned. These 3 factors are thus necessary and sufficient for cumulative route improvements to emerge, even in the absence of sophisticated communication or thought.

Restricted antennal movement impacts the tandem running dynamics in a ponerine ant.

BACKGROUND: Tandem running is a recruitment method found in some species of ants where one ant follows another ant to reach a destination having maintained a physical contact with its antennae, throughout the journey. It is considered that the exchange of information regarding the destination among the nestmates happened during the process of tandem running. We examined the impact of restricting antennal movement on tandem running by using Diacamma indicum, a tandem-running ponerine ant by following 480 tandem runs across 9 treatment colonies and comparing it with 10 control relocating colonies. RESULT: Though all the 19 colonies relocated successfully, treatment colonies took significantly longer time to do so. Restricted antennal movement did not influence the ability to become tandem leaders, initiate tandem runs or the work organization significantly. However, antennae-restricted ants performed fewer tandem runs and took significantly longer time. Followers with single or both antennae-restriction performed significantly higher number of interruptions and the alignment between the leader and follower was impacted as antenna-restricted followers subtended a greater angle and walked more to the side of the leader as compared to the control followers. CONCLUSION: This study showed unhindered movement of the followers' antennae is important for tandem-running ants. In the next step, to gain a comprehensive understanding of this recruitment method, it is essential to individually delineate different sensory modalities.

A new approach to geostatistical synthesis of historical records reveals capuchin spatial responses to climate and demographic change.

Recent proliferation of GPS technology has transformed animal movement research. Yet, time-series data from this recent technology rarely span beyond a decade, constraining longitudinal research. Long-term field sites hold valuable historic animal location records, including hand-drawn maps and semantic descriptions. Here, we introduce a generalised workflow for converting such records into reliable location data to estimate home ranges, using 30 years of sleep-site data from 11 white-faced capuchin (Cebus imitator) groups in Costa Rica. Our findings illustrate that historic sleep locations can reliably recover home range size and geometry. We showcase the opportunity our approach presents to resolve open questions that can only be addressed with very long-term data, examining how home ranges are affected by climate cycles and demographic change. We urge researchers to translate historical records into usable movement data before this knowledge is lost; it is essential to understanding how animals are responding to our changing world.

Nest excavators' learning walks in the Australian desert ant Melophorus bagoti.

The Australian red honey ant, Melophorus bagoti, stands out as the most thermophilic ant in Australia, engaging in all outdoor activities during the hottest periods of the day during summer months. This species of desert ants often navigates by means of path integration and learning landmark cues around the nest. In our study, we observed the outdoor activities of M. bagoti workers engaged in nest excavation, the maintenance of the nest structure, primarily by taking excess sand out of the nest. Before undertaking nest excavation, the ants conducted a single exploratory walk. Following their initial learning expedition, these ants then engaged in nest excavation activities. Consistent with previous findings on pre-foraging learning walks, after just one learning walk, the desert ants in our study demonstrated the ability to return home from locations 2 m away from the nest, although not from locations 4 m away. These findings indicate that even for activities like dumping excavated sand within a range of 5-10 cm outside the nest, these ants learn and utilize the visual landmark panorama around the nest.

Home range of three turtle species in Central Yucatan. A comparative study.

Home range is a fundamental characteristic of an animal natural history. The study of home range provides information on the sites where organisms forage for food, find shelter, or locate mates. Home range size and shape can change throughout the lifespan of an organism, during the year, or across seasons, driven by resource availability and the basic needs for each organism. For freshwater and semi-aquatic turtles, home range is greatly affected by water availability, humidity, and temperature throughout the year, nevertheless demographic factors such age and sex are also important determinants of home range size. In this study we estimated home range and dispersal movements for Kinosternon creaseri, Terrapene yucatana, and Rhinoclemmys areolata in a semi-tropical dry forest in central Yucatán. For a two-year period, turtles were surveyed using hoop traps and visual encounters. Twenty-one individuals (5-8 per species) were equipped with radio transmitters to track them across the landscape. Distances between relocations and home range were compared across species seasons, sex, and interactions of these variables. Monthly average movements were positively correlated with rain in the three species studied. Home range of R. areolata was larger than those of K. creaseri and T. yucatana. Home range of the three studied species were larger during the wet season. Home range overlap index within same species individuals was higher during the rainy than dry season, but overall overlap is low between and within species.

Albatrosses employ orientation and routing strategies similar to yacht racers.

The way goal-oriented birds adjust their travel direction and route in response to wind significantly affects their travel costs. This is expected to be particularly pronounced in pelagic seabirds, which utilize a wind-dependent flight style called dynamic soaring. Dynamic soaring seabirds in situations without a definite goal, e.g. searching for prey, are known to preferentially fly with crosswinds or quartering-tailwinds to increase the speed and search area, and reduce travel costs. However, little is known about their reaction to wind when heading to a definite goal, such as homing. Homing tracks of wandering albatrosses (Diomedea exulans) vary from beelines to zigzags, which are similar to those of sailboats. Here, given that both albatrosses and sailboats travel slower in headwinds and tailwinds, we tested whether the time-minimizing strategies used by yacht racers can be compared to the locomotion patterns of wandering albatrosses. We predicted that when the goal is located upwind or downwind, albatrosses should deviate their travel directions from the goal on the mesoscale and increase the number of turns on the macroscale. Both hypotheses were supported by track data from albatrosses and racing yachts in the Southern Ocean confirming that albatrosses qualitatively employ the same strategy as yacht racers. Nevertheless, albatrosses did not strictly minimize their travel time, likely making their flight robust against wind fluctuations to reduce flight costs. Our study provides empirical evidence of tacking in albatrosses and demonstrates that man-made movement strategies provide a new perspective on the laws underlying wildlife movement.

Home range size and habitat selection of owned outdoor domestic cats (Felis catus) in urban southwestern Ontario.

Domestic cats (Felis catus) play a dual role in society as both companion animals and predators. When provided with unsupervised outdoor access, cats can negatively impact native wildlife and create public health and animal welfare challenges. The effective implementation of management strategies, such as buffer zones or curfews, requires an understanding of home range size, the factors that influence their movement, and the types of habitats they use. Here, we used a community/citizen scientist approach to collect movement and habitat use data using GPS collars on owned outdoor cats in the Kitchener-Waterloo-Cambridge-Guelph region, southwestern Ontario, Canada. Mean (± SD) 100% minimum convex polygon home range size was 8 ± 8 ha (range: 0.34-38 ha) and was positively associated with road density but not with intrinsic factors such as boldness, sex, or age. With regards to habitat selection, cats used greenspaces, roads, and agricultural land less often than predicted but strongly selected for impervious surfaces (urban areas other than greenspaces or roads). Our results suggest that wildlife near buildings and residential areas are likely at the greatest risk of cat predation and that a buffer size of 840 m would be needed to restrict cats from entering areas of conservation concern.

The role of memory-based movements in the formation of animal home ranges.

Most animals live in spatially-constrained home ranges. The prevalence of this space-use pattern in nature suggests that general biological mechanisms are likely to be responsible for their occurrence. Individual-based models of animal movement in both theoretical and empirical settings have demonstrated that the revisitation of familiar areas through memory can lead to the formation of stable home ranges. Here, we formulate a deterministic, mechanistic home range model that includes the interplay between a bi-component memory and resource preference, and evaluate resulting patterns of space-use. We show that a bi-component memory process can lead to the formation of stable home ranges and control its size, with greater spatial memory capabilities being associated with larger home range size. The interplay between memory and resource preferences gives rise to a continuum of space-use patterns-from spatially-restricted movements into a home range that is influenced by local resource heterogeneity, to diffusive-like movements dependent on larger-scale resource distributions, such as in nomadism. Future work could take advantage of this model formulation to evaluate the role of memory in shaping individual performance in response to varying spatio-temporal resource patterns.

Changes in movement patterns in relation to sun conditions and spatial scales in wild western gorillas.

For most primates living in tropical forests, food resources occur in patchworks of different habitats that vary seasonally in quality and quantity. Efficient navigation (i.e., spatial memory-based orientation) towards profitable food patches should enhance their foraging success. The mechanisms underpinning primate navigating ability remain nonetheless mostly unknown. Using GPS long-term tracking (596 days) of one group of wild western lowland gorillas (Gorilla gorilla gorilla), we investigated their ability to navigate at long distances, and tested for how the sun was used to navigate at any scale by improving landmark visibility and/or by acting as a compass. Long episodic movements ending at a distant swamp, a unique place in the home range where gorillas could find mineral-rich aquatic plants, were straighter and faster than their everyday foraging movements relying on spatial memory. This suggests intentional targeting of the swamp based on long-distance navigation skills, which can thus be efficient over a couple of kilometres. Interestingly, for both long-distance movements towards the swamp and everyday foraging movements, gorillas moved straighter under sunlight conditions even under a dense vegetation cover. By contrast, movement straightness was not markedly different when the sun elevation was low (the sun azimuth then being potentially usable as a compass) or high (so providing no directional information) and the sky was clear or overcast. This suggests that gorillas navigate their home range by relying on visual place recognition but do not use the sun azimuth as a compass. Like humans, who rely heavily on vision to navigate, gorillas should benefit from better lighting to help them identify landmarks as they move through shady forests. This study uncovers a neglected aspect of primate navigation. Spatial memory and vision might have played an important role in the evolutionary success of diurnal primate lineages.

The avian olfactory system and hippocampus: Complementary roles in the olfactory and visual guidance of homing pigeon navigation.

The homing pigeon is the foundational model species used to investigate the neural control of avian navigation. The olfactory system is critically involved in implementing the so-called olfactory map, used to locate position relative to home from unfamiliar locations. The hippocampal formation supports a complementary navigational system based on familiar visual landmarks. Insight into the neural control of pigeon navigation has been revolutionised by GPS-tracking technology, which has been crucial for both detailing the critical role of environmental odours for navigation over unfamiliar areas as well as offering unprecedented insight into the role of the hippocampal formation in visual landscape/landmark-based navigation, including a possible, unexpected role in visual-spatial perception.