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Black-tailed prairie dog, Cynomys ludovicianus (Sciuridae), metapopulation response to novel sourced conspecific signals

Aggregation of territorial individuals within a species can be facilitated via conspecific signals, wherein settlement implies habitat suitability, ease of resource acquisition and/or increased predator detection. The black-tailed prairie dog is a colonial small mammal with alarm vocalizations that confer benefits via group vigilance against predators and increased foraging time. Although prairie dog alarm calls are relatively well understood, the information embedded in their jump-yip call, which includes both a distinct cry and a bodily gesture, remains less clear. We evaluated prairie dog behaviour in response to conspecific acoustic signals using playbacks of alarm and jump-yip calls at 26 sites in northeastern Wyoming, U.S.A. Recorded calls from an isolated colony were broadcast to a mean of five individuals per site, and behavioural responses were compared against uninfluenced behaviour and a control playback of ambient sounds. The alarm playback caused prairie dogs to increase vigilance 122% and decrease foraging time 23%, demonstrating prairie dogs will shift behaviour based on signals from individuals of an unfamiliar colony. However, the alarm call playback reduced frequency of the jump-yip behaviour only at colonies nearest the recording source. The jump-yip playback caused unfamiliar prairie dogs to display 339% more jump-yips than uninfluenced behaviour. The jump-yip playback did not alter recipients' foraging or vigilance behaviours relative to control treatments, suggesting that although prairie dogs can understand and reciprocate an unfamiliar, single modality signal, they may not shift other behaviours based on this stimulus. As such, the purpose and benefits of the jump-yip call remain unclear. Playback efficacy also had a nonlinear relationship with distance from recording source. Our work improves understanding of communication at the metapopulation level, examines the potential role of the jump-yip and provides insights for how conspecific signals might be used as a management tool.

Wyoming

Divergent migration within lake sturgeon (Acipenser fulvescens) populations: Multiple distinct patterns exist across an unrestricted migration corridor

Population structure, distribution, abundance, and dispersal arguably underpin the entire field of animal ecology, with consequences for regional species persistence, and provision of ecosystem services. Divergent migration behaviours among individuals or among populations is an important aspect of the ecology of highly-mobile animals, allowing populations to exploit spatially- or temporally-distributed food and space resources. This study investigated the spatial ecology of lake sturgeon ( Acipenser fulvescens ) within the barrier free Huron-Erie Corridor (HEC), which connects Lake Huron and Lake Erie of the North American Laurentian Great Lakes. Over six years (2011 – 2016), movements of 268 lake sturgeon in the HEC were continuously monitored across the Great Lakes using acoustic telemetry (10 yr battery life acoustic transmitters). Five distinct migration behaviours were identified with hierarchical cluster analysis, based on the phenology and duration of river and lake use. Lake sturgeon in the HEC were found to contain a high level of intraspecific divergent migration, including partial migration with the existence of residents. Specific behaviours included year-round river residency and multiple lake-migrant behaviours that involved movements between lakes and rivers. Over 85% of individuals were assign to migration behaviours as movements were consistently repeated over the study, which suggested migration behaviours were consistent and persistent in lake sturgeon. Differential use of specific rivers or lakes by acoustic-tagged lake sturgeon further subdivided individuals into 14 “contingents” (spatiotemporally segregated subgroups). Contingents associated with one river (Detroit or St. Clair) were rarely detected in the other river, which confirmed that lake sturgeon in the Detroit and St. Clair represent two semi-independent populations that could require separate management consideration for their conservation. The distribution of migration behaviours did not vary between populations, sexes, body size, or among release locations, which indicated that intrapopulation variability in migratory behaviour is a general feature of the spatial ecology of lake sturgeon in un-fragmented landscapes.

Michigan, Ontario

A framework for understanding semi-permeable barrier effects on migratory ungulates

1. Impermeable barriers to migration can greatly constrain the set of possible routes and ranges used by migrating animals. For ungulates, however, many forms of development are semi-permeable, and making informed management decisions about their potential impacts to the persistence of migration routes is difficult because our knowledge of how semi-permeable barriers affect migratory behaviour and function is limited. 2. Here, we propose a general framework to advance the understanding of barrier effects on ungulate migration by emphasizing the need to (i) quantify potential barriers in terms that allow behavioural thresholds to be considered, (ii) identify and measure behavioural responses to semi-permeable barriers and (iii) consider the functional attributes of the migratory landscape (e.g. stopovers) and how the benefits of migration might be reduced by behavioural changes. 3. We used global position system (GPS) data collected from two subpopulations of mule deer Odocoileus hemionus to evaluate how different levels of gas development influenced migratory behaviour, including movement rates and stopover use at the individual level, and intensity of use and width of migration route at the population level. We then characterized the functional landscape of migration routes as either stopover habitat or movement corridors and examined how the observed behavioural changes affected the functionality of the migration route in terms of stopover use. 4. We found migratory behaviour to vary with development intensity. Our results suggest that mule deer can migrate through moderate levels of development without any noticeable effects on migratory behaviour. However, in areas with more intensive development, animals often detoured from established routes, increased their rate of movement and reduced stopover use, while the overall use and width of migration routes decreased. 5. Synthesis and applications . In contrast to impermeable barriers that impede animal movement, semi-permeable barriers allow animals to maintain connectivity between their seasonal ranges. Our results identify the mechanisms (e.g. detouring, increased movement rates, reduced stopover use) by which semi-permeable barriers affect the functionality of ungulate migration routes and emphasize that the management of semi-permeable barriers may play a key role in the conservation of migratory ungulate populations.

Journal of Applied Ecology

Short-term responses to a human-altered landscape do not affect fat dynamics of a migratory ungulate

According to risk-sensitive foraging theory, animals should make foraging decisions that balance nutritional costs and gains to promote fitness. Human disturbance is a form of perceived risk that can prompt avoidance of risky habitat over acquisition of food. Consequently, behavioural responses to perceived risk could induce nutritional costs. Population declines often coincide with increases in human disturbance, which likely is associated with direct and indirect habitat loss. Nevertheless, behavioural and physiological responses to perceived risks associated with human disturbance could be an added nutritional deficit with population-level repercussions. Using GPS-collar data from three populations of migratory mule deer Odocoileus hemionus exposed to a gradient of established industrial energy development on winter ranges where direct and indirect habitat loss were well documented, we evaluated whether exposure and behavioural responses to human disturbance alter changes in nutritional condition (i.e. fat reserves) over winter. Although animals exhibited behaviours indicative of perceived risk of human disturbance, such as increased movement rates and avoidance of infrastructure, exposure and behavioural responses to human disturbance had little to no measurable effect on fat loss over a winter. Instead, catabolism of fat reserves occurred primarily as a function of the amount of fat animals had entering winter, suggesting that, in the short term, animals were able to mitigate the energetic costs of perceived risk of human disturbance over winter. Animals, however, did not appear to overcome persistent food limitations, and animals with less food availability lost more fat over winter. Our findings heed caution in using short-term behavioural and physiological responses to inform long-term nutritional consequences of human disturbance. Although animals appear to mitigate the energetic costs of perceived risk, food limitations exacerbated by broader-scale avoidance of food near human disturbance may be the primary pathway causing the frequently observed population declines following human disturbance to pristine landscapes.

Wyoming

Long-term video surveillance and automated analyses reveal arousal patterns in groups of hibernating bats

Understanding natural behaviours is essential to determining how animals deal with new threats (e.g. emerging diseases). However, natural behaviours of animals with cryptic lifestyles, like hibernating bats, are often poorly characterized. White-nose syndrome (WNS) is an unprecedented disease threatening multiple species of hibernating bats, and pathogen-induced changes to host behaviour may contribute to mortality. To better understand the behaviours of hibernating bats and how they might relate to WNS, we developed new ways of studying hibernation across entire seasons. We used thermal-imaging video surveillance cameras to observe little brown bats ( Myotis lucifugus ) and Indiana bats ( M. sodalis ) in two caves over multiple winters. We developed new, sharable software to test for autocorrelation and periodicity of arousal signals in recorded video. We processed 740 days (17,760 hr) of video at a rate of >1,000 hr of video imagery in less than 1 hr using a desktop computer with sufficient resolution to detect increases in arousals during midwinter in both species and clear signals of daily arousal periodicity in infected M. sodalis . Our unexpected finding of periodic synchronous group arousals in hibernating bats demonstrate the potential for video methods and suggest some bats may have innate behavioural strategies for coping with WNS. Surveillance video and accessible analysis software make it now practical to investigate long-term behaviours of hibernating bats and other hard-to-study animals.

Methods in Ecology and Evolution

Use of mammal manure by nesting burrowing owls: a test of four functional hypotheses

Animals have evolved an impressive array of behavioural traits to avoid depredation. Olfactory camouflage of conspicuous odours is a strategy to avoid depredation that has been implicated only in a few species of birds. Burrowing owls, Athene cunicularia, routinely collect dried manure from mammals and scatter it in their nest chamber, in the tunnel leading to their nest and at the entrance to their nesting burrow. This unusual behaviour was thought to reduce nest depredation by concealing the scent of adults and juveniles, but a recent study suggests that manure functions to attract arthropod prey. However, burrowing owls routinely scatter other materials in the same way that they scatter manure, and this fact seems to be at odds with both of these hypotheses. Thus, we examined the function of this behaviour by testing four alternative hypotheses. We found no support for the widely cited olfactory-camouflage hypothesis (manure did not lower the probability of depredation), or for the mate-attraction hypothesis (males collected manure after, not before, pair formation). Predictions of the burrow-occupied hypothesis (manure indicates occupancy to conspecifics and thereby reduces agonistic interactions) were supported, but results were not statistically significant. Our results also supported several predictions of the prey-attraction hypothesis. Pitfall traps at sampling sites with manure collected more arthropod biomass (of taxa common in the diet of burrowing owls) than pitfall traps at sampling sites without manure. Scattering behaviour of burrowing owls appears to function to attract arthropod prey, but may also signal occupancy of a burrow to conspecifics. ?? 2006 The Association for the Study of Animal Behaviour.

Animal Behaviour

Changes in vocal repertoire of the Hawaiian crow, Corvus hawaiiensis,from past wild to current captive populations

For most avian species, social behaviou r is critically important for survival and reproductive success. Many social behaviours in birds are culturally transmitted, and as bird populations decline across the globe, important elements of these behaviours may be lost. The Hawaiian crow or 'alalā, Corvus hawaiiensis , is a socially complex avian species that is currently extinct in the wild. As in other oscine passerines, vocalizations in the 'alalā may be culturally transmitted. We compared the vocal repertoire of three of the last four wild 'alalā pairs from the early 1990s to three current captive pairs on the Island of Hawai'i to determine how acoustic behaviour has been affected by changes in their social and physical environment. Over 18 h of recordings from wild breeding pairs were analysed and compared with 44 h from captive breeding pairs. Calls were placed into five putative behavioural categories: (1) alarm, (2) territorial broadcast, (3) aggression, (4) submission and (5) courtship. There was little difference in the overall number and diversity of call types among wild versus captive birds. However, the repertoire was significantly different. Territorial broadcast calls, common components of the wild repertoire, were absent from the captive repertoire. In addition, wild birds had twice the number of alarm calls and had a higher call rate than captive birds. Our results show how socially learned behaviours may change over relatively short periods for an entire species. Understanding how the vocal repertoire and the functional context of vocalizations change may provide useful information for ongoing efforts to reintroduce the 'alalā into the wild.

Hawai'i

Performance comparisons for artificially propagated and wild Pacific lamprey juveniles and larvae

Artificially propagated Pacific lamprey ( Entosphenus tridentatus ) are produced for restoration and for use in dam passage studies to reduce the demand for wild fish. Such uses require that animals are representative of their wild counterparts. Previous work indicated that this is true for Pacific lamprey larvae and juveniles reared in the hatchery with respect to the length of sustained swimming. However, more subtle differences in behaviour and performance that lamprey need to survive have not been assessed. In this study, artificially propagated and wild fish were compared in laboratory tests under no-flow conditions to examine light avoidance, burrowing speed, burst swim speed, volitional routine swim speed and time to come to rest. Most larvae burrowed in less than a minute, and we found highly significant differences ( p < 0.001) between artificially propagated and wild larvae burrowing times, a critical escape behaviour. This could have implications for studies of larval entrainment at irrigation diversion canals or in turbine boils at dams. Interestingly, all of the wild juveniles tested came to rest quickly after introduction to the chamber (1.5 min), while artificially propagated lamprey swam robotically near the surface and 48% did not come to rest in the first 10 min (median time to rest = 9.5 min). In contrast, wild juveniles quickly (median = 1.47 min) sought areas near the tank bottom and attached strongly with their oral disc. Such behavioural differences could have important survival consequences for artificially propagated lamprey as they approach turbine intakes, bypass screens and irrigation diversion headgates. This study highlights the need to conduct behavioural assays that examine subtleties of fish behaviour that can be missed with traditional swim tunnel comparisons.

Aquaculture, Fish and Fisheries

Snowshoe hares display limited phenotypic plasticity to mismatch in seasonal camouflage

As duration of snow cover decreases owing to climate change, species undergoing seasonal colour moults can become colour mismatched with their background. The immediate adaptive solution to this mismatch is phenotypic plasticity, either in phenology of seasonal colour moults or in behaviours that reduce mismatch or its consequences. We observed nearly 200 snowshoe hares across a wide range of snow conditions and two study sites in Montana, USA, and found minimal plasticity in response to mismatch between coat colour and background. We found that moult phenology varied between study sites, likely due to differences in photoperiod and climate, but was largely fixed within study sites with only minimal plasticity to snow conditions during the spring white-to-brown moult. We also found no evidence that hares modify their behaviour in response to colour mismatch. Hiding and fleeing behaviours and resting spot preference of hares were more affected by variables related to season, site and concealment by vegetation, than by colour mismatch. We conclude that plasticity in moult phenology and behaviours in snowshoe hares is insufficient for adaptation to camouflage mismatch, suggesting that any future adaptation to climate change will require natural selection on moult phenology or behaviour.

Proceedings of the Royal Society B: Biological Sci

On the interface between cultural transmission, phenotypic diversity, demography and the conservation of migratory ungulates

Recent evidence indicates that green-wave surfing behaviour in ungulates and the migrations that stem from this behaviour are socially learned, culturally transmitted across generations and become more efficient via cumulative cultural evolution. But given a lack of corroborative evidence, whether ungulate migration is a cultural phenomenon remains a hypothesis deserving of further testing. In this opinion piece, we summarize the role memory and social learning play in the green-wave surfing that underlies ungulate migration, and when combined with the natural history of ungulates, we argue that the most likely mechanism for maintenance of ungulate migration is animal culture. We further our argument by providing a synopsis of processes that promote diversification of migratory behaviour and link these processes to their emergent ecological patterns, which are common in nature but have not historically been considered as potential cultural phenomena. The notion that diverse portfolios of migratory behaviour may buffer populations from environmental change emerges from this synthesis but requires empirical testing. Finally, we contend that, because the migratory behaviour of ungulates stems largely from cultural transmission as opposed to a genetic programme, the diversity of observed migratory strategies represents ‘culturally significant units’ deserving of the same conservation effort afforded to evolutionarily significant units.

Wyoming

Threading the needle: How humans influence predator–prey spatiotemporal interactions in a multiple‐predator system

Perceived predation risk and the resulting antipredator behaviour varies across space, time and predator identity. Communities with multiple predators that interact and differ in their use of space, time of activity and hunting mode create a complex landscape for prey to avoid predation. Anthropogenic presence and disturbance have the potential to shift interactions among predators and prey and the where and when encounters occur. We examined how white-tailed deer Odocoileus virginianus fawn spatiotemporal antipredator behaviour differed along an anthropogenic disturbance gradient that had black bears Ursus americanus , coyotes Canis latrans , bobcats Lynx rufus and humans present. We quantified (a) spatial co-occurrence in species distributions, (b) temporal overlap across the diel cycle and (c) spatiotemporal associations between humans, bears, coyotes, bobcats, adult male deer and fawns. We also examined how deer vigilance behaviour changed across the anthropogenic disturbance gradient and survey duration. Anthropogenic disturbance influenced spatiotemporal co-occurrence across multiple scales, often increasing spatiotemporal overlap among species. In general, species’ spatial co-occurrence was neutral or positive in anthropogenically disturbed environments. Bears and fawns, coyotes and adult male deer, and bobcats and fawns all had higher temporal overlap in the agriculture-development matrix sites. In addition, factors that influenced deer vigilance (e.g. distance to forest edge and predator relative abundance) in the agriculture-development matrix sites did not in the forest matrix site. By taking into account the different antipredator behaviours that can be detected and the different scales these behaviours might occur, we were able to gain a more comprehensive picture of how humans reduce available niche space for wildlife, creating the neutral and positive spatiotemporal associations between species that studies have been seeing in more disturbed areas.

Pennsylvania

Behavioral state-dependent habitat selection and implications for animal translocations

Post-release monitoring of translocated animals is often used to inform future translocation protocols. Quantifying habitat selection of translocated individuals may help identify features that characterize good settlement habitat and thus inform the choice of future release sites. However, translocated animals often undergo post-release behavioural modification, and their habitat selection may vary depending on the underlying behavioural state. To investigate this, we analysed behavioural state-dependent habitat selection in female greater sage-grouse Centrocercus urophasianus translocated from Wyoming to North Dakota, USA, using Hidden Markov Models combined with Integrated Step Selection Analysis. We segmented individual trajectories into behavioural phases corresponding to an exploratory state, characterized by broad and directed movements, and a restricted state, characterized by short and tortuous movements. Then, we quantified habitat selection in each state while accounting for seasonality and individual reproductive status. While in the exploratory state, sage-grouse exhibited natal habitat preference induction by selecting for high sagebrush cover, which is typical of their natal area in Wyoming but not of the release area in North Dakota. In the restricted state, sage-grouse selected for gentle topography and also adjusted their habitat selection to constraints imposed by seasonality and reproductive needs by selecting for high herbaceous cover during brood rearing. Synthesis and applications . Habitat selection of translocated sage-grouse differed between the post-release exploration and the settlement phase. Features selected after settling, not during exploration, are likely indicative of suitable settlement habitat. Our results suggest that areas characterized by gentle topography and high herbaceous cover are well-suited as release sites for sage-grouse translocated during the summer, especially brood-rearing females, and that sagebrush cover may not be a critical factor in determining the appropriateness of release sites for sage-grouse in North Dakota. Our findings highlight the need to consider behaviour when using habitat selection estimates to inform the choice of future release sites.

North Dakota, Wyoming

Factors influencing Cinnamon Teal nest attendance patterns

Patterns of nest attendance in birds result from complex behaviours and influence the success of reproductive events. Incubation behaviours vary based on individual body condition, energy requirements and environmental factors. We assessed nest attendance patterns in Cinnamon Teal Spatula cyanoptera breeding in the San Luis Valley of Colorado in 2016–2017 using trail and video cameras to observe behaviours throughout incubation. We evaluated the effect of temporal, life-history and environmental covariates on the frequency and duration of incubation recesses as well as the incubation constancy. There was considerable model uncertainty among the models used to evaluate recess frequency. Recess duration varied according to the interaction between nest age and a quadratic effect of time of day, with hens on older nests taking longer recesses in the afternoon and hens on nests earlier in incubation taking longer recesses in the morning and evening. Incubation constancy decreased with higher ambient temperatures in the study area. This study provides evidence that Cinnamon Teal modify their behaviour during incubation according to the age of the nest and the time of day. These results improve our knowledge of Cinnamon Teal breeding ecology and shed light on the behaviours that fast-lived species may use to cope with environmental factors during nesting.

Colorado

Phenotypic divergence of secondary sexual traits among sage grouse, Centrocercus urophasianus , populations

S age grouse, Centrocercus urophasianus , in an isolated montane basin near Gunnison, Colorado differ in several morphological and behavioural traits from conspecifics studied in other areas of the species' range. Both sexes in Gunnison are smaller than sage grouse elsewhere, and males possess differences in feather morphology as well. The mating behaviour of male sage grouse in three populations was examined to determine whether male strut displays of Gunnison sage grouse were behaviourally distinct. Behavioural analyses revealed Gunnison males perform strut displays at a slower rate than males in the two other sage grouse populations sampled. In addition, Gunnison males' strut displays contain unique visual and acoustical aspects. The most distinguishing attributes of Gunnison sage grouse were male secondary sexual characteristics including traits that correlate with mating success in other populations. Thus, phenotypic differences observed in the Gunnison population represent a divergence in expression of traits that are likely to be influenced by sexual selection. Recent models of speciation suggest that species characterized by intense sexual selection, such as those with lek mating systems, have the potential for rapid inter-populational divergence in male traits and female preferences leading to speciation.

Colorado

A new approach to the study of relationship quality in dolphins: framework and preliminary results

Proximity and synchronous behaviours from surface observations have been used to measure association patterns within and between dolphin dyads. To facilitate an investigation of relationship quality in dolphins, we applied a method used for chimpanzees and ravens that examined three main components to describe relationships: value, security, and compatibility. Using pilot data from a long-term study of two study populations for this preliminary assessment, these three components were extracted from PCA analysis of eight behavioural variables with more than 80% variance accounted for in both study groups. Only pair swim position differed between groups. Although value, security, and compatibility are abstract terms, each is based on behaviours identified as important in dolphin social life, at least for these two populations. Examining relationship quality in dolphins with a method used to illustrate dyadic differences for chimpanzees and ravens allows for a quantitative, comparative assessment of sociality across disparate taxa. Although these three species are diverse in their anatomies and in their social habitats (e.g., aquatic, terrestrial, aerial), they may well share the basic societal building blocks in the factors affecting how relationships are formed. We discuss how an examination of these behavioural variables facilitates understanding relationship quality in dolphins, as well as how dolphin relationships fit into the context of social animals’ society.

Bimini Islands, Great Bahama Bank, Roatan Island

Potentially exploitable supercritical geothermal resources in the ductile crust

The hypothesis that the brittle–ductile transition (BDT) drastically reduces permeability implies that potentially exploitable geothermal resources (permeability >10 −16 m 2 ) consisting of supercritical water could occur only in rocks with unusually high transition temperatures such as basalt. However, tensile fracturing is possible even in ductile rocks, and some permeability–depth relations proposed for the continental crust show no drastic permeability reduction at the BDT. Here we present experimental results suggesting that the BDT is not the first-order control on rock permeability, and that potentially exploitable resources may occur in rocks with much lower BDT temperatures, such as the granitic rocks that comprise the bulk of the continental crust. We find that permeability behaviour for fractured granite samples at 350–500 °C under effective confining stress is characterized by a transition from a weakly stress-dependent and reversible behaviour to a strongly stress-dependent and irreversible behaviour at a specific, temperature-dependent effective confining stress level. This transition is induced by onset of plastic normal deformation of the fracture surface (elastic–plastic transition) and, importantly, causes no ‘jump’ in the permeability. Empirical equations for this permeability behaviour suggest that potentially exploitable resources exceeding 450 °C may form at depths of 2–6 km even in the nominally ductile crust.

Nature Geoscience

Birds not in flight: Using camera traps to observe ground use of birds at a wind-energy facility

Context: Camera trapping is increasingly used to collect information on wildlife occurrence and behaviour remotely. Not only does the technique provide insights into habitat use by species of interest, it also gathers information on non-target species. Aims: We implemented ground-based camera trapping to investigate the behaviours of ground-dwelling birds, a technique that has largely been unutilised for studying birds, especially in wind-energy facilities. Methods: We used camera traps to monitor activities of Agassiz’s desert tortoises ( Gopherus agassizii ) at their self-constructed burrows in a wind-energy facility near Palm Springs, California, USA. While doing so, we collected data on numerous burrow commensals, including birds. Key results: Monitoring from late spring to mid-autumn in one year showed regular use of tortoise burrows and the immediate area by 12 species of birds, especially passerines. The most abundant species, as indicated by the number of photographs, but not necessarily individuals, was the rock wren ( Salpinctes obsoletus ), with a total of 1499 events. Birds appeared to use the interior or proximate vicinity of burrows for gathering nesting material, displaying, feeding, dust bathing and other activities. Of the bird species observed, 10 are known to be occasional casualties of turbine-blade strikes. The minimum known-age of a burrow had a positive relationship with bird counts. Conclusions: Using camera traps focused at ground level can be a useful tool in avian conservation efforts because it is an effective technique for measuring bird presence, activity and behaviour in altered habitats such as wind farms, especially for those species that are low flyers or ground dwellers. Implications: Acquiring data over the long term by using ground-based monitoring with camera traps could add to our understanding of avian behaviour and habitat use in relation to wind-energy infrastructure and operations, and help determine the vulnerability of avifauna that utilise the area.

California

Parasite manipulation of brain monoamines in California killifish (Fundulus parvipinnis) by the trematode Euhaplorchis californiensis

California killifish (Fundulus parvipinnis) infected with the brain-encysting trematode Euhaplorchis californiensis display conspicuous swimming behaviours rendering them more susceptible to predation by avian final hosts. Heavily infected killifish grow and reproduce normally, despite having thousands of cysts inside their braincases. This suggests that E. californiensis affects only specific locomotory behaviours. We hypothesised that changes in the serotonin and dopamine metabolism, essential for controlling locomotion and arousal may underlie this behaviour modification. We employed micropunch dissection and HPLC to analyse monoamine and monoamine metabolite concentrations in the brain regions of uninfected and experimentally infected fish. The parasites exerted density-dependent changes in monoaminergic activity distinct from those exhibited by fish subjected to stress. Specifically, E. californiensis inhibited a normally occurring, stress-induced elevation of serotonergic metabolism in the raphae nuclei. This effect was particularly evident in the experimentally infected fish, whose low-density infections were concentrated on the brainstem. Furthermore, high E. californiensis density was associated with increased dopaminergic activity in the hypothalamus and decreased serotonergic activity in the hippocampus. In conclusion, the altered monoaminergic metabolism may explain behavioural differences leading to increased predation of the infected killifish by their final host predators. ?? 2008 The Royal Society.

Proceedings of the Royal Society B: Biological Sci