ESS analysis of mechanistic models for territoriality: the value of scent marks in spatial resource partitioning

In this paper elements of game theory are used to analyse a spatially explicit home range model for interacting wolf packs. The model consists of a system of nonlinear partial differential equations whose parameters reflect the movement behavior of individuals within each pack and whose solutions describe the patterns of space-use by each pack. By modifying the behavioral parameters, packs adjust their patterns of movement so as to maximize their reproductive output. This involves a tradeoff between maximizing prey intake and minimizing conflict with neighbors. Evolutionarily stable choices of the behavioral parameters yields territories that are immune to invasion by groups with alternate behaviors.     

Differences in winter activity,courtship, and social behavior of two captive family groups of Mexican wolves (Canis lupus baileyi)

The purpose of this study was to determine differences in activity patterns and social behavior of two groups of endangered Mexican wolves maintained at two quite different facilities and to determine some of the variables that should be considered when making specific behavioral comparisons among wolves in this binational captive breeding program. Quantitative measurements of an Activity Index and social behaviors were obtained for three individuals in each pack. Within each age/sex category, activity, aggression, and play were more frequent in the pack at a zoo facility, compared to the pack at a field station facility. Frequency of courtship interactions and scent marking were significantly higher in the field station pack. The packs were similar in the frequency of active submission, but differed significantly in the pattern of this behavior. Given the large number of interacting variables and small number of individuals in this study, we recommend caution in generalizing results to other packs or facilities. Zoo Biol 16:435–443, 1997. © 1997 Wiley-Liss, Inc.     

Habitat structure and fish predation: effects on invertebrate colonisation and breakdown of stream leaf packs

1. We investigated the effects of two features of leaf‐pack habitat structure (i.e. mass of a leaf pack and surface area of leaves comprising a leaf pack) and fish predation on colonisation of shredders and leaf breakdown rates in a coldwater stream. Packs were constructed of red maple (Acer rubrum) leaves. 2. A 2 × 3 × 3 factorial experiment was used to manipulate fish predation (exclusion and control cage), leaf‐pack mass (1, 3 and 5 g dry mass) and leaf surface area (small: approx. 17.9 cm², medium: approx. 34.6 cm², large: approx. 65.6 cm²). Exclusion cages had mesh on all sides, whereas control cages lacked mesh on two sides to provide access to fish. 3. Common shredders were Gammarus pseudolimnaeus, Pycnopsyche and Lepidostoma. Shredder biomass per leaf pack increased with the mass of a leaf pack (P < 0.001), but biomass per unit mass of leaf pack did not differ with leaf‐pack mass (P = 0.506). Shredder densities did not respond to the exclusion of fish (P > 0.7) or leaf surface area (P > 0.7), and interactions among treatment factors were not significant (P > 0.2). 4. Breakdown rates were lower for leaf packs comprised of small leaves (P < 0.001) and leaf packs with high mass (P = 0.001). Excluding fish did not significantly affect leaf breakdown rates (P = 0.293), and interactions among treatment factors were not significant (P > 0.3). Breakdown rates were highest when packs consisted of few leaves (i.e. leaf packs with large leaves and low mass) and were colonised by many shredders. 5. Fish predation was not an important factor controlling shredder densities in leaf packs over the spatiotemporal scale of our experiment. Nevertheless, we found shredder colonisation was proportional to leaf‐pack mass and breakdown rates were affected by leaf‐pack size (i.e. number of leaves in a pack). We suspect that fragmentation is the primary mechanism causing the breakdown rates to be dependent on leaf‐pack size.     

The process of a wolf pack splitting in Białowieża Primeval Forest,Poland

In 1998, the pack of 7 wolvesCanis lupus Linnaeus, 1758, radio-tracked in Białowieża Primeval Forest, East Poland, split into 2 packs (2 and 5 wolves), when an 8-year-old alpha female ceased breeding. The two sister-packs subdivided their original territory, but their ranges overlapped extensively (49%) for one year after the split, except for May-June, when both new packs reared pups. We propose that food related factors could have been the ultimate cause of splitting of a large pack. In European temperate forests, pack size of 5–6 wolves is optimal for the consumption of the red deerCervus elaphus.     

Steepness of thermal gradient is essential to obtain a unified view of thermotaxis in C. elegans

One of the adaptive behaviors of animals in their environment is thermotaxis, by which they migrate toward a preferred temperature. This sensorimotor integration is accomplished by choosing one of two behaviors depending on the surrounding temperature, namely thermophilic or cryophilic movement. Caenorhabditis elegans exhibits thermotaxis and its migration behavior has been analyzed experimentally at both the population and individual levels. However, some experimental data are inconsistent especially for thermophilic movement, which is expected to be observed in lower than favorable temperatures. There are no experimental analyzes that find thermophilic tendencies in the individual behavior of worms, despite multiple reports supporting thermophilic movement of the population. Although theoretical methods have been used to study thermotaxis of C. elegans, no mathematical model provides a consistent explanation for this discrepancy. Here we develop a simple biased random walk model, which describes population behavior, but which is based on the results of individual assays. Our model can integrate all previous experiments without any contradiction. We regenerate all the population patterns reported in past studies and give a consistent explanation for the conflicting results. Our results suggest that thermophilic movement is observed, even in individual movements, when the thermal gradient is sufficiently slight. On the contrary, thermophilic movement disappears when the thermal gradient is too steep. The thermal gradient is thus essential for a comprehensive understanding of the experimental studies of thermotaxis in C. elegans. Our model provides insight into an integrative understanding of the neural activity and thermotactic behavior in C. elegans.     

Unexplained patterns of grey wolf Canis lupus natal dispersal

Natal dispersal (movement from the site of birth to the site of reproduction) is a pervasive but highly varied characteristic of life forms. Thus, understanding it in any species informs many aspects of biology, but studying it in most species is difficult. In the grey wolf Canis lupus, natal dispersal has been well studied. Maturing members of both sexes generally leave their natal packs, pair with opposite-sex dispersers from other packs, near or far, select a territory, and produce their own offspring. However, three movement patterns of some natal-dispersing wolves remain unexplained: 1) long-distance dispersal when potential mates seem nearby, 2) round-trip travels from their natal packs for varying periods and distances, also called extraterritorial movements, and often not resulting in pairing, and 3) coincidental dispersal by individual wolves from a given area in the same basic directions and over the same long distances. This perspective article documents and discusses these unexplained dispersal patterns, suggests possible explanations, and calls for additional research to understand them more clearly.     

Ecology of leaf pack macroinvertebrate communities in streams of the Fraser River Basin, British Columbia

1. To characterise geographic and small scale variation in the structure of macroinvertebrate communities in stream leaf packs, we collected one to three natural leaf pack communities from 119 reference streams in the Fraser River Basin and quantified their variability and correlation with aspects of the stream environment at several scales. We also sampled leaf packs in 19 test streams in the same geographic area exposed to stressors (nine logged, seven farmed, three mined catchments) to evaluate the leaf pack community as a tool for bioassessment. 2. There was substantial variation in the composition of invertebrate communities in leaf packs among reference streams of the Fraser River Basin. Capnia and Zapada (stoneflies), Baetis and Ephemerella (mayflies) and Tvetnia (midge) were the most common taxa found in the leaf packs. There were three types of assemblages identified by non‐metric multidimensional scaling; Capnia, Baetis and Ephemerella communities. 3. Leaf pack communities from the 19 test streams were plotted on a non‐metric multidimensional scaling ordination of the reference communities, and 14 of 19 sites fell outside the 80% confidence ellipse of the reference sites, including eight of nine logged, four of seven farmed and one of three mined catchments. Most of these streams plotted on the ordination near the Ephemerella reference communities. Reference stream communities had a similar number of genera per leaf pack (12.0) and genera per site (18.7) as the test streams (12.6 genera per leaf pack and 18.7 genera per site). Among the test sites, the farmed catchments had higher genera per leaf pack (17.8) and genera per site (21.9) than either the logged (11.5 genera per leaf pack; 19.9 genera per site) or mined (3.4 genera per leaf pack; 7.7 genera per site) catchments. 4. Heterogeneity of leaf pack communities within a site decreased as the number of genera found at the site increased. This was determined by allometric regression of the number of genera found at a site on the maximum number of genera possible, given the average number found per leaf pack. 5. There was a significant relationship between the composition of the leaf pack invertebrate community and stream geography (latitude, longitude, altitude, stream order). Canonical correspondence analysis showed differences among ‘big river’, ‘mountain stream’ and ‘southern’ communities. 6. There was no relationship between the composition of the leaf pack invertebrate community and stream channel and flow characteristics (bank dimensions, flow, slope). There was a significant relationship between the composition of the leaf pack invertebrate community and water quality of the stream (oxygen, nitrogen, phosphorus, conductivity, pH, temperature). ‘Cold, oxygen rich water’ communities were distinguishable from communities in streams with warmer, lower oxygen concentration. ‘High nutrient water’ communities were also distinct from communities in low nutrient streams. There was no relationship between the composition of the leaf pack invertebrate community and the nature of the leaf pack itself (i.e. morphology, decomposition, coniferous needle content). 7. Invertebrate communities in leaf packs show substantial, interpretable variation among reference streams. They are sensitive to human stressors at a landscape scale such as forestry and agriculture. Their diversity and composition varies at different spatial scales in a way that is at least partially explained by the environment of the stream and its catchment area.     

Male-biased Adult Sex Ratios and their Significance for Cooperative Breeding in Dhole,Cuon alpinus,Packs

Two free-ranging packs of dholes (Asiatic wild dog, Cuon alpinus) were monitored for a period of 6 yr (Sep. 1990-Sep. 1996) in the Mudumalai sanctuary, southern India. Demographic data on age structure, litter-size, sex ratio and age and sex specific dispersal were collected. Behavioural data on social interactions and reproductive behaviour among pack members were obtained to determine the frequencies of dominant and subordinate behaviours shown by male and female pack members and a measure of each male's reproductive access to females. Behaviours displayed by pack members at dens were recorded to determine whether any age- or sex-specific role specialization existed during pup care. Tenures for dominant males and females within the pack were calculated to ascertain the rate of breeding vacancies occurring within packs. Approximate levels of genetic relatedness within packs were determined by studying pedigrees. In most years one study pack had a male-biased adult sex ratio. This was caused by almost two-fold higher dispersal of adult females over adult males. A considerable variance existed in the percentage of sub-adults dispersing from the two packs. Differences existed in the frequencies of dominant and subordinate behaviours shown by males. For males, dominance ranks and ranks based on submissive behaviours were not correlated with frequencies of reproductive behaviours. Subordinate males also displayed reproductive behaviours. In packs, dominant males had lower tenures than dominant females indicating that among males breeding vacancies arose more quickly. The litter size was found to be negatively correlated with the age of the breeding female. There were no significant differences across individuals of varying age or sex classes in the display of pup care behaviours. Significant differences did exist among individual adults. Genetic relatedness among packs tended to vary temporally as a consequence of possible mating by subordinate animals and immigration of new males into the pack. In conclusion, it appears that males delay dispersal and cooperate within their natal packs because of the variety of reproductive strategies they could pursue within. A combination of ecological constraints and the difficulties of achieving breeding status within non-natal packs may make early dispersal and independent breeding less beneficial.     

Bioturbation by a dominant detritivore in a headwater stream: litter excavation and effects on community structure

Bioturbation can affect community structure by influencing resource distribution and habitat heterogeneity. Bioturbation by detritivores in small headwater streams could affect community structure by reintroducing buried detrital resources into the food web and could also affect the distribution of various taxa on detritus. We evaluated the ability of the caddisfly Pycnopsyche gentilis to uncover experimentally buried leaves in a headwater stream. Packs of leaves were placed in enclosures and covered with a known volume of sediment. We added 0, 3 or 6 large Pycnopsyche to the enclosures which were permeable to most other invertebrate taxa. Leaf packs were sampled after 23 days and leaf pack mass, the amount of sediment covering the leaf packs, and macro‐ and microinvertebrate densities on leaf packs were quantified. There was a significant negative relationship between Pycnopsyche density and leaf pack mass. Pycnopsyche also reduced the volume of sediment covering leaf packs. Pycnopsyche had complex effects on the abundance of invertebrate taxa associated with the leaves. Some taxa exhibited their highest abundance in the 3 Pycnopsyche treatment while others exhibited non‐significant increases as Pycnopsyche density increased. These results suggest that the beneficial effects of Pycnopsyche (e.g. uncovering leaves which increases the availability of habitat and food) outweigh any negative effects (e.g. disturbance, encounter competition) of the caddisfly when it is present at lower densities. However, the negative impacts of Pycnopsyche appear to outweigh the positive effects via sediment removal at higher caddisfly densities for some taxa. Our results suggest that bioturbating organisms in streams have the potential to reintroduce organic matter to detrital food webs and affect the distribution and abundance of benthic taxa associated with organic matter.     

Movements and habitat use in gravid and nongravid female garter snakes (Colubridae: Thamnophis)

Two dhole ( Cuon alpinus ) packs were monitored in Mudumalai Sanctuary, southern India, during 1989-93 to look at population dynamics, movement pattern, and foraging strategy and their inter-relationship with the maintenance of social groups. Pack size fluctuated substantially (4–18 and 4–25 in the two packs) owing to dispersal and demographic factors such as females not breeding in a given year. Both packs killed a much higher proportion of chital ( Axis axis ) and sambar ( Cervus unicolor ) fawns (2 and 54.2km² for the two packs) was not correlated with pack size. Pack movement from one resource patch (consisting of resting sites and aggregations of prey species) to another was not random or based on factors such as inter-patch distance or relative prey densities. There was no difference in mean residence time of the pack across the four resource patches; the pack moved across these in a sequential manner in one direction. We conclude that dholes live in groups not because of any advantages accruing from enhanced group sizes through increased per capita yield of food, but as a consequence of the dispersion of resources.     

Estimation of Successful Breeding Pairs for Wolves in the Northern Rocky Mountains,USA

Abstract: Under the Endangered Species Act, documenting recovery and federally mandated population levels of wolves (Canis lupus) in the Northern Rocky Mountains (NRM) requires monitoring wolf packs that successfully recruit young. United States Fish and Wildlife Service regulations define successful breeding pairs as packs estimated to contain an adult male and female, accompanied by ≥2 pups on 31 December of a given year. Monitoring successful breeding pairs will become more difficult following proposed delisting of NRM wolves; alternatives to historically intensive methods, appropriate to the different ecological and regulatory context following delisting, are required. Because pack size is easier to monitor than pack composition, we estimated probability a pack would contain a successful breeding pair based on its size for wolf populations inhabiting 6 areas in the NRM. We also evaluated the extent to which differences in demography of wolves and levels of human-caused mortality among the areas influenced the probability of packs of different sizes would contain successful breeding pairs. Probability curves differed among analysis areas, depending primarily on levels of human-caused mortality, secondarily on annual population growth rate, and little on annual population density. Probabilities that packs contained successful breeding pairs were more uniformly distributed across pack sizes in areas with low levels of human mortality and stable populations. Large packs in areas with high levels of human-caused mortality and high annual growth rates had relatively high probabilities of containing breeding pairs whereas those for small packs were relatively low. Our approach can be used by managers to estimate number of successful breeding pairs in a population where number of packs and their sizes are known. Following delisting of NRM wolves, human-caused mortality is likely to increase, resulting in more small packs with low probabilities of containing breeding pairs. Differing contributions of packs to wolf population growth based on their size suggests monitoring successful breeding pairs will provide more accurate insights into population dynamics of wolves than will monitoring number of packs or individuals only.     

Population dynamics and food habits of the banded mongoose

A population of approximately 100 banded mongooses living in six packs was studied for over 2 years in Rwenzori National Park, Uganda. The packs were relatively stable, cohesive social units composed of approximately equal numbers of males and females and varied in size from six to thirty-five adults and subadults. Breeding was synchronized within the packs with several females producing their litters at approximately the same time. The packs bred up to four times per year. Mortality of animals over 6 months old was approximately 10% annually. Less than 50% of juveniles survived to the age of 3 months. The banded mongoose feeds primarily on invertebrates, particularly millipedes and beetles. Ants, crickets, termites and earwigs are also important elements of the diet. Vertebrate remains were found in 12% of the droppings analysed. Although the pack forages as a unit each individual finds its own food. The mean home range size of five of the packs was 80-4 ha; the two largest packs had ranges exceeding 1 km². The banded mongoose is strictly diurnal typically leaving a den in the hour following sunrise and returning a few minutes before sunset with a prolonged rest during the heat of the day. Dens were in termite mounds (usually in thicket clumps), erosion gullies, aardvark holes and occasionally in man-made structures. Most were used for only a few days and then abandoned but a few preferred dens were occupied for periods up to 54 days and re-used repeatedly. Interactions between packs were aggressive and appeared to function in spacing the packs. The result of an encounter depended upon pack size with larger packs dominating smaller ones; the area of occurrence did not appear to affect the outcome. Mating between packs was observed during aggressive encounters. Group life is of adaptive significance in protecting the individual from predators and in care of the young. The pack bunches around young mongooses when disturbed and also bunches to attack certain predators and competitors. One or more adults remain at the den to guard the young when the pack forages.     

Movements and diving behavior of weaned Weddell seal (Leptonychotes weddellii ) pups

Between 1993 and 1995, the diving behavior and movement patterns of 23 weaned Weddell seal pups (Leptonychotes weddellii) were tracked in the Ross Sea. Antarctica, using satellite-linked time-depth recorders. Regression analyses revealed that for seals of between 8 and 27 weeks old, age was poorly correlated with the dive depth, duration, or frequency. However, changes in dive parameters suggested that Weddell seal pups were attempting to maximize dive time, but the manner in which this was done depended on age and time of day. Movement patterns indicated that most Weddell seal pups left their natal area by the end of February, and traveled north along the Antarctic continent coastline. Several individuals returned to McMurdo Sound, but others were last located more than 400 km from McMurdo. Routes followed suggest that pups can use the pack ice habitat, but prefer to remain closer to the coastline than do adults. Accepted: 21 July 1998     

Spacing and movement in the green porcelain crab Petrolisthes armatus: evidence for male competition and mate guarding

The green porcelain crab, Petrolisthes armatus, is a common invasive species on inter-tidal oyster reefs in the South Atlantic Bight whose behavior is largely unknown. We assessed the effects of the presence of opposite-sex conspecifics on adult crab spacing at a low density to infer potential mate acquisition behaviors. Adult crabs held in all-male, all-female, and mixed-sex treatments were observed twice daily over five days to determine nearest-neighbor (NN) distances and relative activity. Males in the mixed-sex treatment with the shortest NN distances (NN₁) were most commonly (1) closest to females, and, when compared to NN₁ males in the all-male treatment, were (2) closer to their NN and (3) had lower activity levels. Female movement did not differ between treatments. Percent ovigerous females at the end of trials were higher in the mixed-sex treatment, reflecting mating activity, which when observed, involved inter-molt females. Mate guarding, rather than pure-searching behavior, was the mating behavior most consistent with observed spacing and movement patterns.     

Surveying Predicted Rendezvous Sites to Monitor Gray Wolf Populations

ABSTRACT We used rendezvous site locations of wolf (Canis lupus) packs recorded during 1996–2006 to build a predictive model of gray wolf rendezvous site habitat in Idaho, USA. Variables in our best model included green leaf biomass (Normalized Difference Vegetation Index), surface roughness, and profile curvature, indicating that wolves consistently used wet meadow complexes for rendezvous sites. We then used this predictive model to stratify habitat and guide survey efforts designed to document wolf pack distribution and fecundity in 4 study areas in Idaho. We detected all 15 wolf packs (32 wolf pack-yr) and 20 out of 27 (74%) litters of pups by surveying <11% of the total study area. In addition, we were able to obtain detailed observations on wolf packs (e.g., hair and scat samples) once we located their rendezvous sites. Given an expected decrease in the ability of managers to maintain radiocollar contact with all of the wolf packs in the northern Rocky Mountains, rendezvous sites predicted by our model can be the starting point and foundation for targeted sampling and future wolf population monitoring surveys.     

Over-winter decomposition and associated macroinvertebrate communities of three deciduous leaf species in forest streams on the Canadian Boreal Shield

The decomposition of deciduous leaf material provides a critical source of energy to aquatic food webs. Changes to riparian forests through harvesting practices may alter the species composition of deciduous leaf material entering streams. We compared over-winter decomposition of three different riparian leaf species (speckled alder (Alnus incana ssp. rugosa (Du Roi) J. Clausen), white birch (Betula papyrifera Marsh.), and trembling aspen (Populus tremuloides Michx.)) to determine their importance as a food resource for macroinvertebrate communities within Boreal Shield streams in northeastern Ontario, Canada. Leaf pack decomposition of the three leaf species formed a processing continuum throughout winter, where alder and birch leaf packs decomposed at a medium rate (k = 0.0065/day and 0.0053/day, respectively) and aspen leaf packs decomposed more slowly (k = 0.0035/day). Macroinvertebrate community colonization on leaf packs changed through time regardless of leaf species. Alder leaf packs supported higher abundances of macroinvertebrates in the fall while aspen leaf packs supported greater shredder abundances in the following spring. The study shows that leaf diversity may be important for providing a sustained food resource for aquatic macroinvertebrates throughout the relatively long over-winter period in Canadian Boreal Shield streams. Riparian forest management strategies should ensure that deciduous plant species richness is sustained in riparian areas.     

Annelid sperm and fertilization biology

This paper provides data on fine particulate organic matter (FPOM) and macroinvertebrates associated to natural and artificial leaf packs in a small woodland stream (Schlaube, Brandenburg). Macroinvertebrate colonisation and the dynamics of FPOM were studied in oven-dried alder leaf packs, air-dried alder leaf packs and packs with artificial leafshaped substrate exposed in the stream during a 68-day period. The importance of FPOM as a potential food source for macroinvertebrates especially in artificial leaf packs was evaluated. Changes in the quantity as well as in the chemical composition of the accumulating FPOM (>63 and <63 μm) was determined using soluble carbohydrates, proteins and chlorophyll a as parameters of the nutritional quality. Mass loss and the chemical changes of alder leaves during the decompositional process were also described. The loss of soluble carbohydrates due to leaching was more rapid in oven-dried alder leaf packs than in air-dried ones. After 3 days of leaf pack exposure weight loss of oven-dried and air-dried leaf packs was nearly comparable, as the similar decay coefficients, k = 0.0228 (oven-dried leaf packs) and k = 0.0214 (air-dried leaf packs), respectively, show. The amount of FPOM per unit leaf area constantly increased in artificial packs, although it remained below that of alder leaf packs at all sampling dates. The nutritional quality of FPOM <63 μm was constantly greater than that of FPOM >63 μm and decreased in both size-fractions with length of exposure. Referring to leaf area the abundance of macroinvertebrates continually increased in all packs till the end of exposure, whereas the numbers in artificial packs remained below that in alder leaf packs. The taxonomic composition of all treatments was very similar with Gammarus pulex being the most abundant taxon in all packs until day 42, while afterwards the caddis fly genus Hydropsyche gained in importance. The amphipod Gammarus pulex in general did not show a preference for air-dried alder leaf packs compared to oven-dried alder leaf and artificial packs. Corresponding dynamics of macroinvertebrate colonisation and FPOM content in artificial packs support the hypothesis that FPOM functions not only as an important food source for macroinvertebrates including gammarideans but also as a control mechanism of macroinvertebrate abundance in stream habitats. Even if the accumulation of FPOM and drifting macroinvertebrates might be influenced by the same abiotic factor (e.g. by reduction in stream velocity inside the packs) it is quite unlikely that only physical properties caused the invertebrates to stay.     

How a simple adaptive foraging strategy can lead to emergent home ranges and increased food intake

Animals often alternate between searching for food locally and moving over larger distances depending on the amount of food they find. This ability to switch between movement modes can have large implications on the fate of individuals and populations, and a mechanism that allows animals to find the optimal balance between alternative movement strategies is therefore selectively advantageous. Recent theory suggests that animals are capable of switching movement mode depending on heterogeneities in the landscape, and that different modes may predominate at different temporal scales. Here we develop a conceptual model that enables animals to use either an area‐concentrated food search behavior or undirected random movements. The model builds on the animals’ ability to remember the profitability and location of previously visited areas. In contrast to classical optimal foraging models, our model does not assume food to be distributed in large, well‐defined patches, and our focus is on animal movement rather than on how animals choose between foraging patches with known locations and value. After parameterizing the fine‐scale movements to resemble those of the harbor porpoise Phocoena phocoena we investigate whether the model is capable of producing emergent home ranges and use pattern‐oriented modeling to evaluate whether it can reproduce the large‐scale movement patterns observed for porpoises in nature. Finally we investigate whether the model enables animals to forage optimally. We found that the model was indeed able to produce either stable home ranges or movement patterns that resembled those of real porpoises. It enabled animals to maximize their food intake when fine‐tuning the memory parameters that controlled the relative contribution of area concentrated and random movements.     

A vector-sum process produces curved aiming paths under rotated visual-motor mappings

Under a 90° rotation of motor space relative to visual space, human two-dimensional aiming movements frequently take the form of smooth arcs such as spirals and semi-circles. A time-independent differential equation explains this tendency in terms of a rotation-induced vector field made up, at each point in the two-dimensional space, of two input vectors. One vector represents a visual error signal and the other represents a motor error signal. A trajectory's instantaneous direction of movement at each point can be described as the resultant of the two vectors. This mathematical formulation incorporates plausible visual-motor mechanisms and, when expressed in polar coordinates, leads to a new method for analyzing the spatial properties of movements (i.e., movement paths). Plots of the angle between the resultant and the target vector () against distance from the target (r, in the polar representation) summarize the arc-shaped movement paths as a simple relation that can be analyzed statistically with respect to properties such as monotonicity. The polar representation is a plausible representation of visually-guided movements, with the visual error vector functioning as an objective function relative to which behavior is optimized. We extend the model and ther, movement path analysis to non-90° rotations, and we find that the model predicts an observed qualitative shift in behavior for rotations greater than 90°. It also predicts qualitatively different path shapes observed under visual-motor reflections.This work was performed while the first author was under the support of Grant IST-8511589 from the National Science Foundation and Grant NCC2-307 from the National Aeronautics and Space Administration     

Effects of wolf pack size and winter conditions on elk mortality

Elk (Cervus canadensis) are high-profile game animals for many states in the western United States, yet over the past several decades some populations have experienced a persistent and broad-scale decline in recruitment. Over this same period, gray wolves (Canis lupus) have become an integral component of many western landscapes and agencies are increasingly challenged to maximize hunting opportunities of ungulates via predator management while simultaneously ensuring wolf conservation. To better understand the implications of predator management on elk populations, we monitored survival of 1,244 adult female elk and 806 6-month-old calves from 29 populations distributed throughout Idaho, USA, from 2004 to 2016. We developed predictive models of mortality that related mortality risk to wolf pack size, winter conditions, and individual-level characteristics. Annual mortality rates (excluding harvest) for adult females and calves were 0.09 and 0.40, respectively. Calf mortality was predicted best with a model that included additive effects of chest girth at time of capture, mean size of surrounding wolf packs, and snow depth. Adult female mortality was predicted best with a model that included female age, mean size of surrounding wolf packs, and snow depth. Based on a sensitivity analysis, chest girth had the largest effect on risk of mortality for calves followed by pack size and snow depth. Other than the effect of senescence in the oldest (>15 yr) individuals, pack size and snow depth had the largest effect on risk of mortality for adult females. We estimated cause-specific mortality and predation was the dominant cause of known-fate mortalities for adult females (35% mountain lion [Puma concolor] and 32% wolf) and calves (45% mountain lion and 28% wolf), whereas malnutrition accounted for 9% and 10% of adult female and calf mortalities, respectively. Wolves preferentially selected smaller calves and older adult females, whereas mountain lions showed little preference for calf size or age class of adult females. Our study indicates managers can increase elk survival by reducing wolf pack sizes on surrounding winter ranges, especially in areas where, or during years when, snow is deep. Additionally, managers interested in improving over-winter calf survival can implement actions to increase the size of calves entering winter by increasing the nutritional quality of summer and early fall forage resources. Although our study was prompted by management questions related to wolves, mountain lions killed more elk than wolves and differences in selection of individual elk indicate mountain lions may have comparably more of an effect on elk population dynamics. Although we were unable to relate changes in mountain lion populations to elk survival in our study, future research should seek a better understanding of multi-predator systems, including how management of one predator affect others and ultimately how these interactions affect elk survival. © 2019 The Wildlife Society