Cercariae, like miracidia, are non-parasitic larval stages implicated in the life cycle of all trematodes for the host-to-host parasite transmission. Almost all cercariae are free-living in the external environment. With a few exceptions (cercariae of Halipegus occidualis (Halipegidae) can live several months, Shostak & Esch, 1990a), cercariae have a short active life during which they do not feed, living on accumulated reserves. Most cercariae encyst as metacercariae in second intermediate hosts which are prey of the definitive host; in certain species, the interruption of the active life is achieved by an encystment in the external environment (or a simple immobile waiting strategy in a few species). In some two-host life cycles, the cercariae develop into adults after penetration (this is the case for various species causing human schistosomiasis). Some cercariae do not leave the mollusc which must then be ingested by the definitive host.

Physiological analyses of the behaviour of several cercarial species which actively find and invade their hosts have revealed very complex sequences of behaviour patterns and responses to very different stimuli from the environment and the host. A result of these physiological studies is that the behaviour patterns of each of the species investigated are surprisingly individual. The behavioural patterns of host-finding of those species analysed in some detail reveal profound adaptations to maximize transmission success. This can be demonstrated for movement patterns during swimming, for responses to environmental conditions such as gravity, light and temperature, for responses to stimuli emanating from the host such as shadows, water turbulence and chemical compounds and especially for the responses after contact with the host. The behaviour patterns can be interpreted as adaptations to: (1) dispersal by leaving the habitat of the snail intermediate host and distribution within the area; (2) long survival by energy saving swimming behaviour, by avoiding responses to inappropriate stimuli, by selecting favourable microhabitats and probably by avoiding predation; (3) finding and invading particular host types by selecting microhabitats frequented by the hosts and responding to sequences of specific stimuli emanating from the hosts.

Host location by parasites can be achieved by either active or passive mechanisms. In spite of their significance, the efficacy of these methods has been little researched. High fecundity in parasites is discussed in terms of the role it plays in dispersal and transmission. Some concepts developed by mainstream behavioural ecologists are outlined and their relevance to parasitology is indicated. ‘Reproductive value’ is recommended as an appropriate measure of the costs and benefits of behavioural acts. Although costs of reproduction have been rarely studied in parasites, they are likely to occur in cosexual insects, nematodes and crustaceans. Experiments using captive hosts and/or in vitro cultivation could help in the construction of realistic optimality models. We suggest that r- and K-selection theory could assist in the study of the evolution of parasite behaviour. We discuss how parasite populations are dispersed and controlled and consider the implications of overdispersion. WTe outline three sources of signals to which parasites may respond and suggest that understanding evolutionary mechanisms and community organisation of parasites and hosts requires evaluation of fundamental behavioural responses to environmental signals. The study of closely related groups of parasites and their hosts may advance our knowledge of the evolution of parasite life cycles and the evolutionary costs and benefits of behavioural acts.

Helminth parasites of vertebrates usually select very specific regions or habitats in their hosts, and this is often preceded by a tortuous migration through various host organs. However, the proximate mechanisms of migration and habitat selection have remained enigmatic despite considerable effort by parasitologists. In this paper, a new approach to studying helminth behaviour in the host is proposed. The core idea is that behaviour strategies must be considered from the perspective of the parasites and their perceptions of their environment. A guiding principle is that the environmental features to which an animal responds, and the actions which are required for responding to the environment, form a fundamental unit of behaviour. Thus, we can deduce an animal's behavioural strategy from the details of its response to environmental signals and from its sensory capabilities. The evidence presented suggests that helminth behaviours in the host often occur as fixed (or modal) action patterns which are usually seen in response to constant, or predictable environmental features. Thus, a working hypothesis is that the mechanisms of physiological and biochemical homeostasis within the host provide an extremely predictable environment for the parasite. Under these conditions, a parasite needs to perceive only small subsets of the total information available from the environment to respond appropriately. Studies on sensory and nervous systems of these organisms are critical to understanding parasite perception, but there are formidable technical obstacles that prevent easy access to parasite nervous systems. Therefore, a multidisciplinary approach, using ideas from parasitology, ecology, evolutionary biology and neuroethology, is considered requisite for reconstructing the parasites' behaviour strategies. It is suggested that future directions should pursue integration of studies on sensory physiology with the behavioural ecology of these organisms.

It is generally believed that the patterns of in-host migration of the infective stages of most invasive macroparasites of man and domestic animals are matters of established fact. In reality, the textbook treatment of this topic, and much of the experimental approach to it, are a legacy of an uncritical past. With few exceptions the true picture concerning particular host/parasite systems is confused by the acceptance of inadequate criteria of proof. In important instances these bogus ‘proofs’ have been generalized within a suspect inductive framework which employs the basic assumption that all migratory parasites that enter the same portal and travel to the same destination do so by the same route. The discussion of such issues serves as a background to the presentation in detail of a more rigorous conceptual and experimental framework.

Hutchinson's (1957) definition of an ecological niche as a multidimensional hypervolume determined by a number of physical and biotic variables is adopted. The number of niche dimensions is very great, but as a working hypothesis it is assumed that a few are sufficient to characterize the niche of a parasite species to a high degree of accuracy. They are host species, microhabitat(s), macrohabitat(s), geographical range, sex and age of host, season, food and hyperparasites. Methods to measure niche width, in particular specificity indices, are discussed, and some examples of niche restriction are described. Proximate and ultimate causes of niche restriction are discussed, mainly using marine parasites as examples. Among proximate causes of one niche dimension, host specificity, are ecological factors restricting exposure to infection to certain host species; host-specific chemical factors that induce hatching, direct infective stages to a host and bring about settlement of a parasite; factors that lead to mortality in or on the wrong host; morphological adaptations that guarantee survival in or on the ‘correct’ host; and availability of suitable hosts. Many factors are likely to be responsible for microhabitat specificity, but have been little studied, except for some physiological and morphological adaptations to particular microhabitats. Macrohabitats and geographical range may be determined by the distribution of intermediate hosts and certain food items, and by a variety of chemical and physical factors. Hosts of different sexes may differ in feeding habits and the composition of the skin, and thus acquire parasites differentially. Hosts of different age may be differentially infected due to accumulation of parasites with age, loss of parasites due to developing resistance (or immunity), and different size and feeding habits. Among ultimate causes of niche restriction and segregation are avoidance of competition, predation and hyperparasites; facilitation of mating; reinforcement of reproductive barriers; and adaptations to environmental complexity. Few studies permit a decision on which factor or factors are responsible in particular cases. Interspecific competition may play a greater role in helminth communities of some host groups than of others, but it seems that, overall, its role has been exaggerated at least for marine parasites. Some ‘classical’ examples of microhabitat segregation explained by interspecific competition can also be explained by reinforcement of reproductive barriers. There is evidence for the importance of facilitation of mating in microhabitat restriction, and the availability of many vacant niches indicates that competition, overall, is not of great importance.

This article considers how specificity patterns are shaped during the course of parasite evolution. Parasites are first and foremost specific to site, or microhabitat; host ranges are far more subject to change than is microhabitat. Specificity results from a number of convergent phenomena starting with habits (microhabitat and feeding styles) of free-living progenitors and the way in which the parasitic association arises (e.g., passive oral contamination as opposed to intrusive entry). These bias the types of interaction parasites have with the host, and, through this, the way specificity develops. Host ecology acts as an external factor affecting specificity and predominates in parasites that interact minimally with the hosts physiological and immune systems. Coevolutionary factors are more important in parasites that feed on host tissues or occur in extraintestinal sites. Here, parasites must present the right cues, and respond appropriately to the host defense system. The ability to generalize these cues and responses across host boundaries may act as a constraint on host range. The functional role of the host in the parasite life history also affects the degree of specificity; thus, parasites may act as host generalists in hosts that act as trophic channels to the final host. The role of competition in determining specificity is difficult to assess. However, competition has been reported to influence microhabitat and host distribution through interactive site selection and/or competitive seclusion.

This paper describes a mathematical model which allows us to compare the data collected from short-term cross-sectional surveys with the population dynamics of host and parasite populations over longer periods of time. The model extends an earlier framework for two parasite species in one host, to one for an arbitrary number of parasite species. We show that the conditions necessary for the coexistence of two parasite species extend to expressions for the coexistence of three or more parasite species. Furthermore, the model suggests that those species which form the ‘core’ of the parasite community are those whose high fecundity and transmission efficiency permit them to colonize hosts readily. In contrast, those species which are classified as ‘satellite’ species of the community are either species with low fecundity, or low transmission efficiencies. This work confirms earlier studies that suggest that increasing degrees of aggregation are crucial in allowing several species of parasites to coexist in the same species of hosts.

The properties of the model are compared with patterns observed in data collected for helminth parasites of Anolis lizards, wood mice and eels. This combined theoretical and empirical approach confirms the importance of the life history strategies of the parasite in determining the abundance of each species in the community. It suggests that studies of parasite community structure have to pay more attention to the strategies pursued by each individual species before interactions between species are considered.

Parasite-induced modifications of host behaviour are known from a wide range of host-parasite associations. In many cases, these behavioural changes are thought to be adaptive and. benefit the parasite by increasing its probability of successful transmission. However, in many cases, energy spent on host manipulation will not be available for other functions, such as growth. These trade-offs suggest that in the absence of other constraints, natural selection will optimize, and not maximize, the influence of parasites on host behaviour. This argument is developed and expanded into theoretical considerations of the evolution of host behaviour manipulation by parasites. Among populations of the same parasite species or among closely-related species, the optimal investment into manipulation, or optimal manipulative effort (ME*), of individual parasites is predicted to increase as (1) typical infrapopulation size decreases, (2) prevalence increases, (3) the longevity of the infected host, or of the parasite in its host, decreases, (4) passive transmission rates decrease, and (5) parasite fecundity decreases. This evolutionary analysis indicates that ecological and life history variables may have played an important role in the evolution of manipulation of host behaviour by parasites.

Parasitism is defined in various ways as an intimate relationship in which one partner, the parasite, lives on or in another, the host, generally at the expense of the latter. Parasitism commonly results in a unique array of host physiological responses and adaptations. Most studies of the physiological effects of parasitism have focused on the pathological consequence of infection and disease. While many physiological changes contribute to pathogenesis, it is now recognized that parasitic infections at sub-clinical levels also produce physiological effects that either ameliorate or may not contribute to the disease process. Moreover, these physiological changes are often manifested by altered host behaviour. Behavioural studies have enabled an ecological- and evolutionary-oriented evaluation of host responses. In this fashion, physiological effects may be assessed as to whether they affect fitness and confer benefit or harm to one or both of the symbionts involved. We briefly examine how these physiological responses, specifically neural, endocrine, neuromodulatory, and immunomodulatory components, may interact to modify host behaviours. We consider the adaptiveness of these responses and how the behavioural patterns elicited may simultaneously appear adaptive for the parasite as well as the host. In addition, we address how parasite-host physiological and behavioural interactions may be altered during the course of parasitism.

Behavioural patterns involved in avoiding, repelling, or removing parasites allow mammalian hosts to defend themselves from an array of parasites that threaten the host's fitness in the natural world. Some examples of behavioural defenses and the presumed target parasites are: grooming to remove ticks, grouping to reduce attack rate of biting flies, fly repelling movements to reduce parasitic flies, and selective grazing to reduce intake of faecal-borne parasites. These behavioural defenses are discussed with regard to effectiveness in controlling the target parasites. Parasites have sometimes evolved behavioural strategies of evading, penetrating or disabling these behavioural defenses. These parasite behavioural strategies, though less studied, are discussed. Also discussed is the possibility that host behavioural patterns that may defend against one parasite may be exploited by a different type of parasite to facilitate its own transmission. The interplay between host defensive strategies, the cost versus the effectiveness of such strategies, and a parasite's evasion or exploitation of such strategies, may be useful in understanding some aspects of host-parasite dynamics in nature.