The Science and Philosophy of a Method for Assessing Environmental Causes

When an environmental impairment has been identified, it becomes necessary to identify the cause so that an appropriate action can be planned. However, causation is difficult to establish—both conceptually and in practice. To ensure that the U.S. Environmental Protection Agency's (USEPA's) method for causal assessment is appropriate and defensible, we reviewed concepts of causation from philosophers, statisticians, epidemiologists, and others. This article summarizes the results of that review and explains how it relates to the USEPA's method. We include a five-step process: (1) identify alternative candidate causes; (2) logically eliminate when possible; (3) diagnose when possible; (4) analyze the strength of evidence for remaining candidate causes; and (5) identify the most likely cause. We also encourage three practices: (1) use a consistent process; (2) do not claim proof of causation; and (3) document the evidence and inferences. This approach allows assessors to identify the most likely cause or, failing that, to reduce the set of possible causes and identify information needs for another iteration of causal assessment.     

Rethinking Causation for Data-intensive Biology: Constraints,Cancellations, and Quantized Organisms

Complex organisms thwart the simple rectilinear causality paradigm of “necessary and sufficient,” with its experimental strategy of “knock down and overexpress.” This Essay organizes the eccentricities of biology into four categories that call for new mathematical approaches; recaps for the biologist the philosopher's recent refinements to the causation concept and the mathematician's computational tools that handle some but not all of the biological eccentricities; and describes overlooked insights that make causal properties of physical hierarchies such as emergence and downward causation straightforward. Reviewing and extrapolating from similar situations in physics, it is suggested that new mathematical tools for causation analysis incorporating feedback, signal cancellation, nonlinear dependencies, physical hierarchies, and fixed constraints rather than instigative changes will reveal unconventional biological behaviors. These include “eigenisms,” organisms that are limited to quantized states; trajectories that steer a system such as an evolving species toward optimal states; and medical control via distributed “sheets” rather than single control points.     

CARE OF CEREBRAL-PALSIED CHILDREN—The Special Role of Pediatricians

Because the cerebral-palsied child is also a growing person whose development should be encouraged along as normal a course as possible, the pediatrician may well take the lead in coordinating orthopedic, psychiatric, educational and vocational services for such children. In El Centro this policy has been followed in a school for handicapped children serving 30 with cerebral palsy, 15 of them pupils at the school. Those with sufficient intelligence and milder physical handicap attend a regular public school, while others are unable to attend even the special school.Emphasis has been placed on working with children who are less seriously affected. As to children with severe neuromuscular and intellectual handicap, the chief effort is to prevent contractures and maintain function. The pediatrician confers monthly with all workers concerned with the child, and maintains liaison with the family physician who treats acute illnesses, including palsy seizures. Special effort has been made to investigate family circumstances such as foreign background which make it more difficult to evaluate the palsied child''s true capabilities.     

Exploring the implications of recreational disturbance on an endangered butterfly using a novel modelling approach

Site and wildlife managers globally are under increasing pressure to implement management strategies that address the negative implications of outdoor recreational activities on wildlife. For many rare and isolated species any anthropogenic activities that cause disturbance could potentially be detrimental to existing populations. Understanding how non-consumptive recreation can influence a species may therefore be critical to its preservation. We developed a novel approach to specifically address this need. Using a combination of field surveys and simulation modelling exercises, we (1) explored the responses of endangered Karner blue butterflies (Lycaeides melissa samuelis) to recreation, (2) assessed whether such responses influenced oviposition rate and/or host plant choice and (3) tested alternative management strategies that could alleviate the negative impacts of recreation. Our field surveys confirmed that Karner blues were sensitive to recreational disturbance. Butterflies flushed at similar speeds and distances from recreationists (2.2 m at 0.17 m/s), as they would from natural threats, such as predators (2.2 m at 0.19 m/s). Incorporating female response parameters into a simulation model revealed that regular disturbance could reduce egg laying potential and significantly restrict host plant choice, which in turn, could impact the butterfly’s population dynamics. However, we established that it was possible to effectively offset the implications of recreational disturbance using our simulation modelling approach. For example, extending Karner blue breeding habitat from trails and other public rights of way has the potential to alleviate such disturbance. Our study demonstrates that the potential impact of recreation on species of conservation concern should not be overlooked.     

Pattern,process, and natural disturbance in vegetation

Natural disturbances have been traditionally defined in terms of major catastrophic events originating in the physical environment and, hence, have been regarded as exogenous agents of vegetation change. Problems with this view are: (1) there is a gradient from minor to major events rather than a uniquely definable set of major catastrophes for each kind of disturbance, and (2) some disturbances are initiated or promoted by the biotic component of the system. Floras are rich in disturbance-adapted species. Disturbances have probably exerted selective pressure in the evolution of species strategies. Heathland cyclic successions and gap-phase dynamics in forests have been viewed as endogenous patterns in vegetation. When death in older individividuals imposes a rhythm on community reproduction, dynamics may indeed be the result of endogenous factors. However, documented cases of senescence in perennial plants are few and many cyclic successions and cases of gap-phase dynamics are initiated by physical factors. Forest dynamics range from those that are the result of individual tree senescence and fall, through those that are the result of blowdown of small groups of healthy trees, to those that are the result of large windstorms which level hectares of forest. The effect of wind ranges from simple pruning of dead plant parts to widespread damage of living trees. Wind speed is probably inversely proportional to occurrence frequency. Disturbances vary continuously. There is a gradient from those community dynamics that are initiated by endogenous factors to those initiated by exogenous factors. Evolution has mediated between species and environment; disturbances are often caused by physical factors but the occurrence and outplay of disturbances may be a function of the state of the community as well. Natural disturbances in North American vegetation are: fire, windstorm, ice storm, ice push on shores, cryogenic soil movement, temperature fluctuation, precipitation variability, alluvial processes, coastal processes, dune movement, saltwater inundation, landslides, lava flows, karst processes, and biotic disturbances. Disturbances vary regionally and within one landscape as a function of topography and other site variables and are characterized by their frequency, predictability, and magnitude. The landscape level is important in assessing disturbance regime. Disturbances and cyclic successions belong to the same class of events—that of recurrent dynamics in vegetation structure—irrespective of cause. Dynamics may result from periodic, abrupt, and catastrophic environmental factors or they may result from an interaction of the changing susceptability of the community and some regular environmental factor. In any case, the dynamics result in heterogeneous landscapes; the species adapted to this heterogeneity are numerous, suggesting their long time importance. The importance of disturbance regime as part of the environmental context of vegetation means that allogenic and autogenic models of vegetation are difficult to apply. Species composition can be seen to be a function of disturbance regime, as well as other environmental variables. Competitive replacement in succession occurs, then, only as disturbances cease to operate and can be viewed as allogenic adjustment to a new disturbance-free environment. Competitive divergence, separation of role, and competition avoidance may, in fact, underlie successional patterns traditionally viewed as the competitive replacement of inferior species by superiorly adapted climax species. The importance of ongoing dynamics is also difficult to reconcile with the concept of climax, founded as it is on the idea of autogenesis within a stable physical environment. Climax composition is relative to disturbance regime. Climax is only arbitrarily distinguished from succession. Climax as an organizing paradigm in plant ecology has obscured the full temporal-spatial dimensions important in understanding the vegetated landscape and the evolution of species which contribute to the landscape patterns. Whittaker’s coenocline concept is accepted with modifications: (1) natural disturbance gradients and Whittaker’s complex gradient are intimately related, (2) temporal variation in the community should be viewed as an added axis of community pattern, and (3) ongoing dynamics have important effects on specificity of species to site relations and the predictability of vegetation patterns. Recent work has suggested an r-K continuum in species strategy. In general, colonizing ability is seen as a trade-off against specialization. Frequent disruption of the community and the creation of open sites seems to result in mixes of species that are fleeting in time and do not repeat in space. Species in such mixes are often tolerant of wide environmental extremes but are compressed into early successional time if disturbance ceases. The composition of such communities is not predictable from site characteristics. Even communities with low disturbance frequency lack complete environmental determinism, and historical events are important in understanding present composition. Communities vary in level of environmental determinism and species differ in niche breadth and degree of site specificity. Management implications of vegetation dynamics are discussed.     

Getting rid of interventions

According to James Woodward’s influential interventionist account of causation, X is a cause of Y iff, roughly, there is a possible intervention on X that changes Y. Woodward requires that interventions be merely logically possible. I will argue for two claims against this modal character of interventions: First, merely logically possible interventions are dispensable for the semantic project of providing an account of the meaning of causal statements. If interventions are indeed dispensable, the interventionist theory collapses into (some sort of) a counterfactual theory of causation. Thus, the interventionist theory is not tenable as a theory of causation in its own right. Second, if one maintains that merely logically possible interventions are indispensable, then interventions with this modal character lead to the fatal result that interventionist counterfactuals are evaluated inadequately. Consequently, interventionists offer an inadequate theory of causation. I suggest that if we are concerned with explicating causal concepts and stating the truth-conditions of causal claims we best get rid of Woodwardian interventions.     

Observations on the Etiology and Therapy of “Brittle” Diabetes

Salient aspects of prolonged metabolic studies on seven excessively labile diabetic patients and a review of the literature concerning causation and therapy of brittle diabetes are presented. Brittleness is redefined as “a syndrome of excessive insulin-sensitivity and ketosis-proneness manifested by extreme and unexplainable short-term and long-term fluctuations in the parameters of the disease”. Evidence on the causation of hyperlability points to dysfunction of plasma-protein transport and of hepatic and peripheral tissue metabolism of insulin. No objectively demonstrable complete and lasting stabilization was possible by means of any antidiabetic or adjunctive therapeutic measures. However, achievement of quantitative improvement in the accuracy of regulation of diabetes and moderation in deviations from the acceptable range of parameters were feasible. To this end, therapy recommended for everyday use incorporates the following principles found to be most helpful in following the oscillations of the disease on the research ward: flexibility in the plan of therapy; accuracy, especially in timing of therapeutic events; and employment of an insulin program best suited to the patient''s needs and comfort.     

Promoting Sleep in Shiftworkers and Inter Continental Travelers

A number of techniques and treatments can be used to alleviate the sleep disturbance associated with both shiftwork and transmeridian travel. Optimization of the sleeping environment and avoidance of substances such as caffeine and alcohol before sleep are the best initial approach. Timing sleep to coincide with some of the normal sleep period where possible will improve sleep quality in shiftworkers. Similarly, following transmeridian flight, restricting sleep to the nocturnal period in the new time zone will assist adaptation. Hypnotic drugs may be of benefit to alleviate sleep disturbance experienced by shiftworkers or transmeridian travelers. Selection of the most appropriate medication must take into account required duration of action and possible residual effects of the drug on alertness. Hypnotics may be useful, particularly in middle-aged individuals who already have disturbed sleep, on those occasions when poor sleep is anticipated, for example following an eastward flight or after the initial change to night duty. Over-the-counter preparations should be avoided whenever possible unless it is known that they are not associated with residual sequelae. (Chronobiology International, 14(2), 133–143, 1997)     

Bird sensitivity to disturbance as an indicator of forest patch conditions: An issue in environmental assessments

An Environmental Assessment (EA) is one of the steps within the Environmental Impact Assessment process. Birds are often used in EA to help decision makers evaluate potential human impacts from proposed development activities. A “sensitivity to human disturbance” index, created by Parker III et al. (1996) for all Neotropical species, is commonly considered an ecological indicator. However, this parameter was created subjectively and, for most species, there have been no rigorous field test to validate its effectiveness as such. Therefore, in this study, we aim to: (1) evaluate if, at the local scale, birds from forest patches in a human-modified landscape (HML) may differ in sensitivity from Parker's sensitivity classification; (2) evaluate the effectiveness of the species richness value at each sensitivity level as an ecological indicator; (3) gather information on how often and in which manner Parker's classification has been used in EA. To do so, bird sampling was performed in eight forest patches in a HML over one year. Then, we created a local sensitivity to disturbance using information about threat, endemism, spatial distribution and relative abundance of all species in the study area. We found that 37% of the forest birds showed different local sensitivity levels when compared with Parker's classification. Our results show that only the richness of high-sensitivity species from our local classification fitted the ecological indicator assumptions helping the environmental conditions evaluation of the studied patches. We conclude that species richness of each Parker's bird sensitivity levels do not necessarily perform as an ecological indicator at the local scale, and particularly in HML. Nevertheless, Parker's Neotropical bird sensitivity classification was used in 50% of EA we reviewed. In these, 76% assumed that it was an accurate ecological indicator of the local forest conditions for birds. The lack of clear criteria used in Parker's classification allows diverse interpretations by ornithologists, and there is no agreement about the ecological meaning of each sensitivity level and what environmental conditions each level may indicate of. Therefore, the use of Parker's classification in EA may jeopardize accurate interpretations of proposed anthropogenic impacts. Furthermore, because a bird species’ sensitivity often varies between locations, we argue that Parker's generalized classification of bird sensitivity should not be used as an indicator of forest environmental conditions in EA throughout HMLs in Neotropics. Rather, local bird ecological indices should be explored, otherwise, erroneous predictions of the anthropogenic impacts will continue to be common.     

Managing visitor access to seabird colonies: a spatial simulation and empirical observations

Managers of wildlife reserves have a range of tools available to them when considering the best way to provide visitor access while avoiding as many of the negative effects of human disturbance as possible. However, managers lack guidelines as to whether conservation interests are best met by spreading visitors thinly throughout a reserve or by aggregating them in a small area. Here I describe how relationships between disturbance impact and disturbance pressure (the dose–response curve) can be used to address this issue. I generate a spatial simulation of two different models of visitor distribution (one more aggregated than the other) and explicitly model disturbance impact for a variety of dose–response curves. I show that the optimal visitor distribution is likely to depend on the sensitivity of the species and the overall visitor pressure. Importantly, I find that in certain circumstances optimal management can shift from one management option to the other if visitor numbers cross a certain threshold. I use published relationships predicting nesting success of Common Guillemots Uria aalge and Black-legged Kittiwakes Rissa tridactyla to assess optimal management at three nature reserves in Scotland. Optimal management for Guillemots depends on the number of people and the distance between the people and the birds. At sites with high disturbance pressures, management should aim to aggregate visitors in as small an area as possible, whereas at sites with lower disturbance pressure, an even distribution of visitors is favoured. Kittiwake models were not generally accurate, and consequently only site-specific guidelines could be generated, where an even distribution was favoured.     

On models and muddles of heritability

One reason for the astonishing persistence of the IQ myth in the face of overwhelming prior and posterior odds against it may be the unbroken chain of excessive heritability claims for ‘intelligence’, which IQ tests are supposed to ‘measure’. However, if, as some critics insist, ‘intelligence’ is undefined, and Spearman's g is beset with numerous problems, not the least of which is universal rejection of Spearman's model by the data, then how can the heritability of ‘intelligence’ exceed that of milk production of cows and egg production of hens? The thesis of the present review paper is that the answer to this riddle has two parts: (a) the technical basis of heritability claims for human behavior is just as shaky as that of Spearman's g. For example, a once widely used ‘heritability estimate’ turns out to be mathematically invalid, while another such estimate, though mathematically valid, never fits any data; and (b) valid technical criticisms of flawed heritability claims typically are met with stubborn editorial resistance in the main stream journals, which tends to calcify such misinformation.     

Plant intelligence: Why,why not or where?

The concept of plant intelligence, as proposed by Anthony Trewavas, has raised considerable discussion. However, plant intelligence remains loosely defined; often it is either perceived as practically synonymous to Darwinian fitness, or reduced to a mere decorative metaphor. A more strict view can be taken, emphasizing necessary prerequisites such as memory and learning, which requires clarifying the definition of memory itself. To qualify as memories, traces of past events have to be not only stored, but also actively accessed. We propose a criterion for eliminating false candidates of possible plant intelligence phenomena in this stricter sense: an “intelligent” behavior must involve a component that can be approximated by a plausible algorithmic model involving recourse to stored information about past states of the individual or its environment. Re-evaluation of previously presented examples of plant intelligence shows that only some of them pass our test.

“You were hurt?” Kumiko said, looking at the scar.Sally looked down. “Yeah.”“Why didn''t you have it removed?”“Sometimes it''s good to remember.”“Being hurt?”“Being stupid.”—(W. Gibson: Mona Lisa Overdrive)

Key words: intelligence, memory, learning, plant development, mathematical models, plant neurobiology, definition of terms     

The Ethics of Genetic Research of Intelligence

Should research on the possible genetic components of human intelligence be carried out? I first try to provide some general guidelines as to whether any particular piece of research should be undertaken and then consider the specific example of the ethics of genetic research on intelligence. The history of the debate on intelligence does not make one very optimistic that the fruits of such research would be used wisely. However, there are indications that people’s understanding of the nature of inheritance may be improving and it could be that such research might have significant benefits. It is worth remembering than the condition phenylketonuria, a genetic disease in any useful sense of the term, and one that leads to mental retardation (i.e. very low intelligence), is now wholly preventable, and indeed very largely prevented, through environmental intervention.     

A conceptual framework for the spatial analysis of functional trait diversity

Functional trait diversity is a popular tool in modern ecology, mainly used to infer assembly processes and ecosystem functioning. Patterns of functional trait diversity are shaped by ecological processes such as environmental filtering, species interactions and dispersal that are inherently spatial, and different processes may operate at different spatial scales. Adding a spatial dimension to the analysis of functional trait diversity may thus increase our ability to infer community assembly processes and to predict change in assembly processes following disturbance or land‐use change. Richness, evenness and divergence of functional traits are commonly used indices of functional trait diversity that are known to respond differently to large‐scale filters related to environmental heterogeneity and dispersal and fine‐scale filters related to species interactions (competition). Recent developments in spatial statistics make it possible to separately quantify large‐scale patterns (variation in local means) and fine‐scale patterns (variation around local means) by decomposing overall spatial autocorrelation quantified by Moran's coefficient into its positive and negative components using Moran eigenvector maps (MEM). We thus propose to identify the spatial signature of multiple ecological processes that are potentially acting at different spatial scales by contrasting positive and negative components of spatial autocorrelation for each of the three indices of functional trait diversity. We illustrate this approach with a case study from riparian plant communities, where we test the effects of disturbance on spatial patterns of functional trait diversity. The fine‐scale pattern of all three indices was increased in the disturbed versus control habitat, suggesting an increase in local scale competition and an overall increase in unexplained variance in the post‐disturbance versus control community. Further research using simulation modeling should focus on establishing the proposed link between community assembly rules and spatial patterns of functional trait diversity to maximize our ability to infer multiple processes from spatial community structure.     

Procreative beneficence: why we should select the best children

Eugenic selection of embryos is now possible by employing in vitro fertilization (IVF) and preimplantation genetic diagnosis (PGD). While PGD is currently being employed for the purposes of detecting chromosomal abnormalities or inherited genetic abnormalities, it could in principle be used to test any genetic trait such as hair colour or eye colour.
Genetic research is rapidly progressing into the genetic basis of complex traits like intelligence and a gene has been identified for criminal behaviour in one family. Once the decision to have IVF is made, PGD has few 'costs' to couples, and people would be more inclined to use it to select less serious medical traits, such as a lower risk of developing Alzheimer Disease, or even for non-medical traits. PGD has already been used to select embryos of a desired gender in the absence of any history of sex-linked genetic disease.
I will argue that: (1) some non-disease genes affect the likelihood of us leading the best life; (2) we have a reason to use information which is available about such genes in our reproductive decision-making; (3) couples should select embryos or fetuses which are most likely to have the best life, based on available genetic information, including information about non-disease genes. I will also argue that we should allow selection for non-disease genes even if this maintains or increases social inequality. I will focus on genes for intelligence and sex selection.
I will defend a principle which I call Procreative Beneficence: couples (or single reproducers) should select the child, of the possible children they could have, who is expected to have the best life, or at least as good a life as the others, based on the relevant, available information.     

Medical Care of the Alcoholic Patient

Alcoholism is an illness that constitutes a major health problem at all levels of society. The physician should accept his responsibility to prevent it and to care for the alcoholic. If he knows that one of his patients is drinking immoderately, he should warn him of the outlook. A patient''s acquired dependence on alcohol may be overt, or revealed only on examination for organic disease or emotional disturbance. The diagnosis may be accepted reluctantly, or denied despite positive evidence, but the patient should be persuaded to give up drinking. He may require psychiatric help or advice from a social worker. He may be so ill as to require treatment in hospital, and hospitals must recognize the urgency of such admissions. Discharge from hospital does not end treatment, for alcoholism is a chronic disease, requiring long-term planning, persistent follow-up and enduring sympathy by the physician, who must always be as available to his alcoholic patient as he is to his patient with diabetes, epilepsy or cardiac disease.     

Genome-wide association study of handedness excludes simple genetic models

Handedness is a human behavioural phenotype that appears to be congenital, and is often assumed to be inherited, but for which the developmental origin and underlying causation(s) have been elusive. Models of the genetic basis of variation in handedness have been proposed that fit different features of the observed resemblance between relatives, but none has been decisively tested or a corresponding causative locus identified. In this study, we applied data from well-characterised individuals studied at the London Twin Research Unit. Analysis of genome-wide SNP data from 3940 twins failed to identify any locus associated with handedness at a genome-wide level of significance. The most straightforward interpretation of our analyses is that they exclude the simplest formulations of the ‘right-shift'' model of Annett and the ‘dextral/chance'' model of McManus, although more complex modifications of those models are still compatible with our observations. For polygenic effects, our study is inadequately powered to reliably detect alleles with effect sizes corresponding to an odds ratio of 1.2, but should have good power to detect effects at an odds ratio of 2 or more.     

The Relationship between Trait Emotional Intelligence and Interaction with Ostracized Others' Retaliation

Background

Regulation of emotions in others is distinct from other activities related to trait emotional intelligence in that only such behavior can directly change other people''s psychological states. Although emotional intelligence has generally been associated with prosociality, emotionally intelligent people may manipulate others'' behaviors to suit their own interests using high-level capabilities to read and manage the emotions of others. This study investigated how trait emotional intelligence was related to interacting with ostracized others who attempt retaliation.

Method

We experimentally manipulated whether two people were simultaneously ostracized or not by using an online ball-tossing game called Cyberball. Eighty university students participated in Cyberball for manipulating ostracism and a “recommendation game,” a variation of the ultimatum game for assessing how to interact with others who attempt retaliation, with four participants. After the recommendation game, participants rated their intention to retaliate during the game.

Results

People with higher interpersonal emotional intelligence were more likely to recommend that the ostracized other should inhibit retaliation and maximize additional rewards when they have a weaker intention to retaliate. However, they were more likely to recommend that the ostracized other should retaliate against the ostracizers when they have a stronger intention to retaliate.

Conclusion

This is the first laboratory study that empirically reveals that people with high interpersonal emotional intelligence influence others'' emotions based on their own goals contrary to the general view. Trait emotional intelligence itself is neither positive nor negative, but it can facilitate interpersonal behaviors for achieving goals. Our study offers valuable contributions for the refinement of the trait emotional intelligence concept in the respect of its social function.     

Prolonged recovery time after eruptive disturbance of a deep-sea hydrothermal vent community

Deep-sea hydrothermal vents are associated with seafloor tectonic and magmatic activity, and the communities living there are subject to disturbance. Eruptions can be frequent and catastrophic, raising questions about how these communities persist and maintain regional biodiversity. Prior studies of frequently disturbed vents have led to suggestions that faunal recovery can occur within 2–4 years. We use an unprecedented long-term (11-year) series of colonization data following a catastrophic 2006 seafloor eruption on the East Pacific Rise to show that faunal successional changes continue beyond a decade following the disturbance. Species composition at nine months post-eruption was conspicuously different than the pre-eruption ‘baseline'' state, which had been characterized in 1998 (85 months after disturbance by the previous 1991 eruption). By 96 months post-eruption, species composition was approaching the pre-eruption state, but continued to change up through to the end of our measurements at 135 months, indicating that the ‘baseline'' state was not a climax community. The strong variation observed in species composition across environmental gradients and successional stages highlights the importance of long-term, distributed sampling in order to understand the consequences of disturbance for maintenance of a diverse regional species pool. This perspective is critical for characterizing the resilience of vent species to both natural disturbance and human impacts such as deep-sea mining.     

Escape Tactics and Alarm Responses in Badgers Meles meles: A Field Experiment

A typical badger (Meles meles) territory contains a primary burrow or 'main sett' plus several secondary burrows or 'outlier setts'. The main aim of our study was to test the hypothesis that outlier setts are used as emergency refuges, by subjecting foraging radio-collared badgers to three levels of experimental disturbance (low, moderate and high). In addition, we recorded the occurrence of potential alarm signals. With low-level disturbance, badgers usually returned to the main sett even when this was further away than the nearest outlier sett; with high-level disturbance they always took refuge in the nearest outlier; and with moderate-level disturbance they showed no clear preference. We conclude that outlier setts do act as emergency refuges, but only when an animal is badly frightened. Possible alarm signals (pilo-erection, head-flagging, snorting and growling) occurred mainly during moderate or high-level disturbance but even then they were relatively infrequent. Signals were no more likely to be emitted when conspecifics were near by than when the signaller was alone, and when conspecifics were present they rarely reacted either to the flight of the disturbed animal or to any signals that it emitted. We conclude that such alarm signals as do occur constitute threats directed towards the predator rather than warnings for the benefit of conspecifics.