Morality: An Evolutionary Account Krebs, Dennis L.
Perspectives on psychological science,
05/2008, Letnik:
3, Številka:
3
Journal Article
Recenzirano
Refinements in Darwin's theory of the origin of a moral sense create a framework equipped to organize and integrate contemporary theory and research on morality. Morality originated in deferential, ...cooperative, and altruistic "social instincts," or decision-making strategies, that enabled early humans to maximize their gains from social living and resolve their conflicts of interest in adaptive ways. Moral judgments, moral norms, and conscience originated from strategic interactions among members of groups who experienced confluences and conflicts of interest. Moral argumentation buttressed by moral reasoning is equipped to generate universal and impartial moral standards. Moral beliefs and standards are products of automatic and controlled information-processing and decision-making mechanisms. To understand how people make moral decisions, we must understand how early evolved mechanisms in the old brain and recently evolved mechanisms in the new brain are activated and how they interact. Understanding what a sense of morality is for helps us understand what it is.
Snowshoe hare cycles are one of the most prominent phenomena in ecology. Experimental studies point to predation as the dominant driving factor, but previous experiments combining food ...supplementation and predator removal produced unexplained multiplicative effects on density. We examined the potential interactive effects of food limitation and predation in causing hare cycles using an individual‐based food‐supplementation experiment over‐winter across three cycle phases that naturally varied in predation risk. Supplementation doubled over‐winter survival with the largest effects occurring in the late increase phase. Although the proximate cause of mortality was predation, supplemented hares significantly decreased foraging time and selected for conifer habitat, potentially reducing their predation risk. Supplemented hares also lost less body mass which resulted in the production of larger leverets. Our results establish a mechanistic link between how foraging time, mass loss and predation risk affect survival and reproduction, potentially driving demographic changes associated with hare cycles.
Snowshoe hare cycles are one of the most notable phenomena in ecology. Through an individual‐based food supplementation experiment, we found strong evidence that both food limitation and predation interactively drive hare cycles through changes in behaviour and condition.
In this article, the authors evaluate L.
Kohlberg's (1984)
cognitive-developmental approach to morality, find it wanting, and introduce a more pragmatic approach. They review research designed to ...evaluate Kohlberg's model, describe how they revised the model to accommodate discrepant findings, and explain why they concluded that it is poorly equipped to account for the ways in which people make moral decisions in their everyday lives. The authors outline in 11 propositions a framework for a new approach that is more attentive to the purposes that people use morality to achieve. People make moral judgments and engage in moral behaviors to induce themselves and others to uphold systems of cooperative exchange that help them achieve their goals and advance their interests.
Motion‐sensitive cameras are commonly used to monitor wildlife occupancy rates; however, few studies have assessed whether data from cameras are correlated with density estimates obtained from more ...traditional labor‐intensive methods such as those based on capture‐mark‐recapture. We used data from a boreal forest community to test whether camera data were correlated with densities estimated from independent monitoring methods. We placed 72 covert cameras in the forest around Lhù'ààn Mân' (Kluane Lake), Yukon, Canada, for 7 years and tracked changes in population densities by camera hit rates. We independently estimated population densities of snowshoe hares (Lepus americanus) and red squirrels (Tamiasciurus hudsonicus) using capture‐mark‐recapture via live trapping, and Canada lynx (Lynx canadensis), coyotes (Canis latrans), and moose (Alces americanus) by snow track transects. Density estimates obtained from conventional aerial surveys were also periodically available for moose. Except for red squirrels, camera hit rates were highly correlated with population density estimates obtained by traditional methods, including across a large range of estimated densities corresponding to cyclic population dynamics in several species. Accordingly, we infer that motion‐sensitive cameras could supplement or replace traditional methods for monitoring key species in boreal forest food webs. Using cameras to monitor population change has several advantages; they require less effort in the field, are non‐invasive compared to live‐trapping, include multiple species at the same time, and rely less on weather than either aerial surveys or snow track transects. Tracking changes across the vast boreal forest is becoming increasingly necessary because of climate and landscape change and our data validate the use of motion‐sensitive cameras to provide a useful quantitative method for state‐of‐the‐environment reporting.
We used data from a Yukon boreal forest community to test whether motion‐sensitive camera data were correlated with density estimated from independent monitoring. Camera hit rates were highly correlated with capture‐mark‐recapture density estimates for snowshoe hares and were highly correlated with snow track transect density estimates for lynx and coyotes. Camera hit rates were not correlated with independent density estimates for red squirrels.
The assumption that activity and foraging are risky for prey underlies many predator-prey theories and has led to the use of predator-prey activity overlap as a proxy of predation risk. However, the ...simultaneous measures of prey and predator activity along with timing of predation required to test this assumption have not been available. Here, we used accelerometry data on snowshoe hares (
) and Canada lynx (
) to determine activity patterns of prey and predators and match these to precise timing of predation. Surprisingly we found that lynx kills of hares were as likely to occur during the day when hares were inactive as at night when hares were active. We also found that activity rates of hares were not related to the chance of predation at daily and weekly scales, whereas lynx activity rates positively affected the diel pattern of lynx predation on hares and their weekly kill rates of hares. Our findings suggest that predator-prey diel activity overlap may not always be a good proxy of predation risk, and highlight a need for examining the link between predation and spatio-temporal behaviour of predator and prey to improve our understanding of how predator-prey behavioural interactions drive predation risk.
Some mammal species inhabiting high-latitude biomes have evolved a seasonal moulting pattern that improves camouflage via white coats in winter and brown coats in summer. In many high-latitude and ...high-altitude areas, the duration and depth of snow cover has been substantially reduced in the last five decades. This reduction in depth and duration of snow cover may create a mismatch between coat colour and colour of the background environment, and potentially reduce the survival rate of species that depend on crypsis. We used long-term (1977-2020) field data and capture-mark-recapture models to test the hypothesis that whiteness of the coat influences winter apparent survival in a cyclic population of snowshoe hares (
) at Kluane, Yukon, Canada. Whiteness of the snowshoe hare coat in autumn declined during this study, and snowshoe hares with a greater proportion of whiteness in their coats in autumn survived better during winter. However, whiteness of the coat in spring did not affect subsequent summer survival. These results are consistent with the hypothesis that the timing of coat colour change in autumn can reduce overwinter survival. Because declines in cyclic snowshoe hare populations are strongly affected by low winter survival, the timing of coat colour change may adversely affect snowshoe hare population dynamics as climate change continues.
Extensive research confirms that environmental stressors like predation risk can profoundly affect animal condition and physiology. However, there is a lack of experimental research assessing the ...suite of physiological responses to risk that may arise under realistic field conditions, leaving a fragmented picture of risk-related physiological change and potential downstream consequences on individuals. We increased predation risk in free-ranging snowshoe hares (Lepus americanus) during two consecutive summers by simulating natural chases using a model predator and monitored hares intensively via radio-telemetry and physiological assays, including measures designed to assess changes in stress physiology and overall condition. Compared to controls, risk-augmented hares had 25.8% higher free plasma cortisol, 15.9% lower cortisol-binding capacity, a greater neutrophil: lymphocyte skew, and a 10.4% increase in glucose. Despite these changes, intra-annual changes in two distinct condition indices, were unaffected by risk exposure. We infer risk-augmented hares compensated for changes in their stress physiology through either compensatory foraging and/or metabolic changes, which allowed them to have comparable condition to controls. Although differences between controls and risk-augmented hares were consistent each year, both groups had heightened stress measures during the second summer, likely reflecting an increase in natural stressors (i.e., predators) in the environment. We show that increased predation risk in free-ranging animals can profoundly alter stress physiology and that compensatory responses may contribute to limiting effects of such changes on condition. Ultimately, our results also highlight the importance of biologically relevant experimental risk manipulations in the wild as a means of assessing physiological responses to natural stressors.
Food availability and temporal variation in predation risk are both important determinants of the magnitude of antipredator responses, but their effects have rarely been examined simultaneously, ...particularly in wild prey. Here, we determine how food availability and long-term predation risk affect antipredator responses to acute predation risk by monitoring the foraging response of free-ranging snowshoe hares (Lepus americanus) to an encounter with a Canada lynx (Lynx canadensis) in Yukon, Canada, over four winters (2015–2016 to 2018–2019). We examined how this response was influenced by natural variation in long-term predation risk (2-month mortality rate of hares) while providing some individuals with supplemental food. On average, snowshoe hares reduced foraging time up to 10 h after coming into close proximity (≤75 m) with lynx, and reduced foraging time an average of 15.28 ± 7.08 min per lynx encounter. Hares tended to respond more strongly when the distance to lynx was shorter. More importantly, the magnitude of hares’ antipredator response to a lynx encounter was affected by the interaction between food-supplementation and long-term predation risk. Food-supplemented hares reduced foraging time more than control hares after a lynx encounter under low long-term risk, but decreased the magnitude of the response as long-term risk increased. In contrast, control hares increased the magnitude of their response as long-term risk increased. Our findings show that food availability and long-term predation risk interactively drive the magnitude of reactive antipredator response to acute predation risk. Determining the factors driving the magnitude of antipredator responses would contribute to a better understanding of the indirect effects of predators on prey populations.
Coyotes and lynx are the two most important mammalian predators of snowshoe hares throughout much of the boreal forest. Populations of hares cycle in abundance, with peaks in density occurring every ...8-11 yr, and experimental results suggest that predation is a necessary factor causing these cycles. We measured the functional responses of coyotes and lynx during a cyclic fluctuation of hare populations in the southwest Yukon, to determine their effect on the cyclic dynamics. We used snow-tracking and radio telemetry to examine changes in the foraging behavior of the predators. Coyotes and lynx both fed mostly on hares during all winters except during cyclic lows, when the main alternative prey of coyotes was voles, and lynx switched to hunting red squirrels. Both predators showed clear functional responses to changes in the densities of hares. Kill rates of hares by coyotes varied from 0.3 to 2.3 hares/d, with the most hares killed one year before the cyclic peak, while those of lynx varied from 0.3 to 1.2 hares/d, with the highest one year after the peak. Maximum kill rates by both predators were greater than their energetic needs. The functional response of coyotes was equally well described by linear and type-2 curves, and that of lynx was well described by a type-2 curve. Kill rates by coyotes were higher during the increase in density of hares than during the cyclic decline, while the reverse was true for lynx. Coyotes killed more hares early in the winter, and cached many of these for later retrieval. Lower densities of hares were associated with longer reactive distances of both predators to hares, but with little apparent change in time spent searching or handling prey. In summary, our data show that the two similarly sized predators differed in their foraging behavior and relative abilities at capturing alternative prey, leading to different patterns in their functional responses to fluctuations in the density of their preferred prey.
The boreal forest, the world’s largest terrestrial biome, is undergoing dramatic changes owing to anthropogenic stressors, including those of climate change. To track terrestrial ecosystem changes ...through space and time, robust monitoring programs are needed that survey a variety of ecosystem constituents. We monitored white spruce (
Picea glauca
) cone crops, berry (
Empetrum nigrum, Shepherdia canadensis
) production, above-ground mushroom abundance, and the abundance of small mammals (
Clethrionomys rutilus
,
Peromyscus maniculatus
), North American red squirrels (
Tamiascirus hudsonicus
), snowshoe hares (
Lepus americanus
), and carnivores (
Lynx canadensis
,
Canis latrans
,
Vulpes vulpes
,
Martes americana
,
Mustela erminea
) across 5 sites in the Yukon, Canada. Monitoring began in 1973 at Lhù’ààn Mân’ (Kluane Lake) and additional protocols were added until a complete sequence was fixed in 2005 at all 5 sites and continued until 2022. White spruce cone counts show mast years at 3–7-year intervals. Ground berries and soapberry counts were highly variable among sites and counts did not correlate among sites or between years for different species. Red-backed voles showed clear 3–4-year cycles at Kluane and probably at the Mayo and Watson Lake sites, but showed only annual cycles in Whitehorse and Faro. Snowshoe hares fluctuated in 9–10-year cycles in a travelling wave, peaking one year earlier at Watson Lake but in synchrony at all other sites, with no clear sign of peak density changing or cyclic attenuation over the last 50 years. Red squirrel numbers at Kluane exhibit marked inter-year variability, driven mainly by episodic white spruce cone crops and predation from Canada lynx and coyotes as hare densities undergo cyclic decline. Snow track counts to index mammalian predators have been conducted on our Kluane and Mayo sites, indicating that lynx numbers rise and fall with a 1–2-year lag at these two sites, tracking the hare cycle. Coyotes and lynx at Kluane peak together following the hare cycle, but coyote counts are also depressed during deep snow years. To summarize, we noted considerable inter-site variability in the population dynamics of many boreal forest ecosystem constituents, but the keystone species (snowshoe hare, Canada lynx) exhibit remarkably similar population trends across the region. We continue to monitor wildlife abundance, cone crops, berry production, and mushroom biomass to determine changes associated with increasing temperature and fluctuating rainfall. The Yukon boreal forest is changing as climate shifts, but the changes are slow, variable across sites, taxa specific, and of uncertain predictability.
