Microhabitat segregation in three rocky intertidal fish species in Portugal: does it reflect interspecific competition?

Recruits of Lipophrys pholis , Coryphoblennius galerita and Gobius cobitis spend the low tide under water in pools, and in the case of G. cobitis also in channels. The pools used by the two blenniids show greater similarity than those used by G. cobitis. Larger juveniles and adults of C. galerita continue to use the same type of pools, which are temporarily abandoned by the males during the breeding season when they use emersed holes and crevices as nests. The much faster growing L. pholis and G. cobitis leave the pools in their first autumn, when they are c. 7 cm long. Whereas G. cobitis >7 cm move to channels and large upper pools, with a smooth substratum, sand and boulders, L. pholis begin to use emersed crevices where they spend low tide often in large groups. When L. pholis or C. galerita were added or removed selectively from pools, their numbers returned to pre-experimental levels after 2 weeks, suggesting that, although intraspecific competition may be strong, interspecific competition was unlikely to explain these results fully.     

Rapid learning of shelter position in an intertidal fish, the shanny Lipophrys pholis L.

The homing ability of an intertidal fish, the shanny Lipophrys pholis , was investigated using two experiments that were based on the shanny's natural propensity to home to a refuge. A displacement experiment demonstrated that the fish were able to accurately locate the previous position of a refuge once the shelter itself had been removed so that it could not be used as a cue to directly signal the goal location. This shows that the shanny can encode information about its familiar surroundings into a spatial map and use this information to home. A second experiment in which the cues internal and external to the experimental tank were put in conflict with one another suggested that the shanny can encode cues that are both intra- and external-tank cues in its representation of space, but that there is individual variation in the type of cues that are used, or memorized.     

A comparison of circatidal rhythmicity and entrainment by hydrostatic pressure cycles in the rock goby, Gobius paganellus L. and the shanny, Lipophrys pholis (L.)

The intertidal teleosts Gobius paganellus and Lipophrys pholis show endogenous circatidal activity rhythms when recorded in constant conditions. Under these conditions, the rhythm of L. pholis is the more precise which may indicate stronger coupling between underlying circalunadian oscillators in this species. In G. pagunellus the inter-oscillator coupling may be weaker and this could enable a more subtle interpretation of tidal fluctuations than in L. pholis . The oscillators may, however, be fundamentally different in the two species; circalunadian in G. paganellus and circatidal in L. pholis .
When exposed to hydrostatic pressure cycles of tidal frequency both species responded pre- dominantly to increasing pressure, which suggests that in the wild they are likely to be most active on the rising tide. Hydrostatic pressure cycles are confirmed as a zeitgeber for both species by the successful entrainment of some individuals. The lack of entrainment of others impIies that additional zeitgebers are required for complete entrainment.     

Experimental manipulation reveals the importance of refuge habitat temperature selected by lizards

Refuges provide shelter from predators, and protection from exposure to the elements, as well as other fitness benefits to animals that use them. In ectotherms, thermal benefits may be a critical aspect of refuges. We investigated microhabitat characteristics of refuges selected by a heliothermic scincid lizard, Carlia rubrigularis, which uses rainforest edges as habitat. We approached lizards in the field, simulating a predator attack, and quantified the refuge type used, and effect of environmental temperatures (air temperature, substrate temperature and refuge substrate temperature) on the amount of time skinks remained in refuges after hiding (emergence time). In respone to our approach, lizards were most likely to flee into leaf litter, rather than into rocks or woody debris, and emergence time was dependent on refuge substrate temperature, and on refuge substrate temperature relative to substrate temperature outside the refuge. Lizards remained for longer periods in warmer refuges, and in refuges that were similar in temperature to outside. We examined lizard refuge choice in response to temperature and substrate type in large, semi‐natural outdoor enclosures. We experimentally manipulated refuge habitat temperature available to lizards, and offered them equal areas of leaf litter, woody debris and rocks. When refuge habitat temperature was unmanipulated, lizards (85%) preferred leaf litter, as they did in the field. However, when we experimentally manipulated the temperature of the leaf litter by shading, most skinks (75%) changed their preferred refuge habitat from leaf litter to woody debris or rocks. These results suggest that temperature is a critical determinant of refuge habitat choice for these diurnal ectotherms, both when fleeing from predators and when selecting daytime retreats.     

Patterns of agonistic behaviour, shelter occupation and habitat preference in juvenile Lipophrys pholis, Coryphoblennius galerita and Gobius cobitis

The patterns of agonistic behaviour and substratum preferences were investigated in captive groups of juveniles of Lipophrys pholis (Blenniidae), Coryphoblennius galerita (Blenniidae) and Gobius cobitis (Gobiidae). In monospecific groups, size was an excellent predictor of the rank attained by each individual. In heterospecific groups, both C. galerita and L. pholis were dominant over G. cobitis of similar size, and C. galerita dominated L. pholis . Although G. cobitis showed higher rates of activity and of attacks per minute, and a higher ratio of attacks over threats, all these measures were depressed in the presence of either blenniid. In both blennies, in monospecific groups, the dominance rank of each fish was a good predictor of the time spent inside shelters. G. cobitis occupied preferentially the sandy substratum both in mono- and heterospecific groups. Both blennies, when in monospecific groups, occupied preferentially the rocky substratum, with C. galerita showing the highest level of selectivity. In nature, C. galerita also showed a marked preference for rocky substrata, while G. cobitis was especially abundant in mixed bottoms. L. pholis occupied an intermediate position. In the blenniids studied, competition for access to shelter may be one major functional consequence of agonistic behaviour in non-reproductive contexts.     

Parasitoid and moth movement from refuge to crop

Refuges may provide habitat and resources which enhance population size of natural enemies of herbivorous pests and add to their colonisation into target crops, potentially increasing effectiveness of biological control. They may also allow for an increase in numbers of herbivorous pests. In addition, these refuges may be attractive so that natural enemies aggregate there, neglecting predation and parasitism of pests in the crop. We investigate the movements of marked Diadegma semiclausum (Hellén) a key larval parasitoid, and the adult stage of the host it attacks, diamondback moth, Plutella xylostella (L.), from an on-farm flowering refuge of Brassica vegetables to the Brassica crop. Our results showed that parasitoids and moths visit the flowering refuge, and that they move from the refuge into the crop before and after disturbing the refuge by ploughing. For D. semiclausum, the pattern of movement from the refuge into the crop was similar for males and females, but more males than females were captured. Ploughing the refuge resulted in increased dispersal by males but not females into the crop. Neither moths nor parasitoids preferentially aggregated around the flowering refuge, instead most often their spatial pattern was random throughout the field indicating that they are quite vagile, quickly moving to distances >100 m. This basic understanding of the movements of these insects has implications beyond this system and is relevant to any agricultural system implementing a refuge to enhance natural enemy effectiveness.     

Physiological stress and refuge behavior by African elephants

Physiological stress responses allow individuals to adapt to changes in their status or surroundings, but chronic exposure to stressors could have detrimental effects. Increased stress hormone secretion leads to short-term escape behavior; however, no studies have assessed the potential of longer-term escape behavior, when individuals are in a chronic physiological state. Such refuge behavior is likely to take two forms, where an individual or population restricts its space use patterns spatially (spatial refuge hypothesis), or alters its use of space temporally (temporal refuge hypothesis). We tested the spatial and temporal refuge hypotheses by comparing space use patterns among three African elephant populations maintaining different fecal glucocorticoid metabolite (FGM) concentrations. In support of the spatial refuge hypothesis, the elephant population that maintained elevated FGM concentrations (iSimangaliso) used 20% less of its reserve than did an elephant population with lower FGM concentrations (Pilanesberg) in a reserve of similar size, and 43% less than elephants in the smaller Phinda reserve. We found mixed support for the temporal refuge hypothesis; home range sizes in the iSimangaliso population did not differ by day compared to nighttime, but elephants used areas within their home ranges differently between day and night. Elephants in all three reserves generally selected forest and woodland habitats over grasslands, but elephants in iSimangaliso selected exotic forest plantations over native habitat types. Our findings suggest that chronic stress is associated with restricted space use and altered habitat preferences that resemble a facultative refuge behavioral response. Elephants can maintain elevated FGM levels for ≥ 6 years following translocation, during which they exhibit refuge behavior that is likely a result of human disturbance and habitat conditions. Wildlife managers planning to translocate animals, or to initiate other management activities that could result in chronic stress responses, should consider the potential for, and consequences of, refuge behavior.     

The influence of predator presence on utilization of artificial seagrass habitats by juvenile walleye pollock,Theragra chalcogramma

Synopsis Behavioral preference for a structured habitat (artificial seagrass) by juvenile walleye pollock,Theragra chalcogramma, was tested in controlled laboratory experiments. We monitored position of fish in 2000 1 tanks with and without artificial seagrass present in one half of the tank. In addition, we exposed walleye pollock to a predator model, assessing their response when a grass plot was available or unavailable as a potential refuge. In the absence of predators, the fish avoided the artificial seagrass, displaying a preference for the open water side of the experimental tanks. In the presence of a predator model, however, juvenile walleye pollock readily entered the artificial seagrass plots. In addition, they often remained in the grass canopy in proximity to the predator instead of moving out of the grass to avoid the predator (when no grass was present they consistently moved to the opposite side of the tank from the predator). The behavioral choices exhibited in this study suggest that juvenile walleye pollock modify habitat selection in response to perceived predation risk, and recognize the structure provided by artificial seagrass as a potential refuge.     

Roe deer at risk: teasing apart habitat selection and landscape constraints in risk exposure at multiple scales

Non‐consumptive effects of predators result from the cost of responses to perceived risk. Prey modulate risk exposure through flexible habitat selection at multiple scales which, in interaction with landscape constraints, determines their use of risky habitats. Identifying the relative contributions of landscape constraints and habitat selection to risk exposure is a critical first step towards a mechanistic understanding of non‐consumptive effects. Here, we provide an integrative multi‐scale study of roe deer spatial responses to variable hunting pressure along a landscape gradient of open habitats and dispersed refuges. Between low‐risk and high‐risk periods, we investigated shifts in 1) home‐range location, 2) probability of using risky habitats (between‐habitat scale) and 3) distance to the nearest refuge (within‐habitat scale). For 2) and 3), we disentangled the contributions of landscape constraints and habitat selection to risky habitat use. We found that when risk was high, roe deer did not shift their home‐range, but generally decreased their use of risky habitats, and sometimes reduced their distance to cover (particularly older animals). There was a functional response in between‐habitat selection, with animals living in more open landscapes responding more than those living in landscapes with more refuges. However, individuals living in more open landscapes avoided open risky habitat less. Finally, we found that among‐individual variation in risk exposure was generally, but not always, minimized by habitat selection across gradients of landscape constraints. To our knowledge, this is the first study simultaneously documenting prey responses to risk at the within‐habitat, between‐habitat and home‐range scales. Our results support the view that between‐habitat selection acts at a higher hierarchical level than within‐habitat selection, and provide a framework for disentangling the contributions of habitat selection and landscape constraints to risk exposure. Selection cannot always compensate for landscape constraints, indicating a need for further investigation of the processes underlying habitat selection.     

Vitellogenin gene expression in the intertidal blenny Lipophrys pholis: a new sentinel species for estrogenic chemical pollution monitoring in the European Atlantic coast?

The presence of estrogenic chemicals (ECs) in the aquatic environment is a growing problem. While most attention was initially given to fresh water and estuarine ecosystems, it is now evident that coastal marine areas are also vulnerable to these pollutants. The use of vitellogenin induction in male fish, a specific biomarker of EC exposure, has been the most widely applied methodology. However, in some occasions, the high mobility and migratory behaviour of common sentinel fish species makes data interpretation difficult. Hence, there is the need to validate new sentinel marine fish species which should display, among other features, a strong homing behaviour. The shanny, Lipophrys pholis, is an intertidal fish that combines many of the required characteristics for a sentinel species: abundance and easy of catch, wide geographical distribution and restricted home range. Thus, in order to evaluate, in the field, the species sensitivity to ECs, L. pholis males were collected at two sites reflecting different degrees of anthropogenic contamination. The vitellogenin II gene (VTGII) was isolated and its liver expression evaluated by RT-PCR in the field samples. A significant induction of gene expression was observed in the specimens collected in the urban area, if compared to the reference site, which suggests exposure to ECs. Moreover, a 21-days laboratory exposure to environmental relevant concentrations of ethinylestradiol (EE2) was also performed. A significant induction of L. pholis VTGII gene in EE2 exposed males was observed suggesting similar sensitivity to that of other marine/estuarine fishes. Even though further validation is currently in progress, the available data indicates that L. pholis is responsive to ECs, thus favouring its future integration in monitoring programmes designed to evaluate the presence of ECs in European marine ecosystems.     

Hunger, light level and body size affect refuge use by post-settlement lingcod Ophiodon elongatus

Using experimental methods, body size, nutritional state (hunger) and light level were investigated as possible determinants of the extent to which juvenile lingcod Ophiodon elongatus used structural refuges. In the laboratory, older, larger individuals (134–162 mm total length, L _T) exhibited a higher propensity for structure use than did smaller juvenile fish (78–103 mm L _T). In fish of both body sizes, increasing hunger levels caused fish to emerge more readily from refuges than satiated individuals, with this effect most prominent for smaller fish. Diel patterns of ambient illumination were simulated in the laboratory, and refuge use by large individuals decreased significantly in response to lower nocturnal light levels, a pattern that was not observed in the smaller size class. These results indicate an ontogenetic shift in refuge use. During the summer, as post-settlement fish grew larger, they tended to increase affinity for structurally complex habitat, and also showed divergent patterns of behaviour in relation to hunger and light level.     

An experimental test of condition-dependent mating behavior and habitat choice by water striders in the wild

Individual condition is expected to be an important determinantof many behaviors, including mating dynamics and habitat choice.In this study we experimentally investigated the linkages betweenindividual condition, habitat use, and mating dynamics in thewild. We manipulated recent feeding history of the water strider,Aquarius remigis, and then quantified the habitat use and matingactivity of males and females. Females could choose from threehabitats (refuge, near shore, and open). On the water surface(open and near-shore habitats), in contrast to refuge, femalescan forage, but they are exposed to predation and sexual harassmentby males. We tested three main hypotheses. First, we predictedthat single females that were fed, relative to those that werenot, would reduce their exposure to predators and male harassmentby increasing their use of refuge. Similarly, we predicted thatfed females that were mating would spend more time in refugethan those that were not fed. Our results support these twopredictions. Fed single and mating females significantly increasedtheir use of refuge. Third, we predicted that mating activity(proportion of time spent mating) of fed females would be reducedrelative to starved females, because of reduced exposure tomales while in refuge, and perhaps because of decreased receptivityto male mating attempts while on the water surface. Mating activityof fed females was about one third that of starved females.The decrease in mating activity could not be accounted for byany change in female receptivity, but could be accounted forby change in habitat use. The decrease in mating activity mayhave resulted from decreases in both mating frequency and matingduration. Our estimates of minimum mating frequency indicatea large and significant decrease, but we were unable to assessmating duration. We found no significant effect of our manipulationon habitat use or mating activity of males.[Behav Ecol 7: 474–479(1996)]     

Habitat separation of semifree-ranging Lemur catta and Lemur fulvus

Habitat use of two semifree-ranging groups of lemurs, Lemur catta and Lemur fulvus, was studied in relation to abiotic, floristic, and structural characteristics of their enclosure. L. catta was very flexible in the use of the habitat and showed a tendency for random utilization. But this species reacted quickly to changes of the abiotic environment. L. fulvus preferred to stay in areas with dense vegetation but good lateral visibility and close to the food plants. Abiotic factors such as solar radiation and ambient temperature influenced L. fulvus to a lesser extent than L. catta. According to these results L. catta could be considered to be adapted to variable environments where it is advantageous to react quickly to changes of abiotic factors. The conservative habitat use and the limited response to abiotic factors of L. fulvus might reflect adaptations of a species living in a stable environment. If these interpretations prove correct, then this study provides another example of animals born and raised in captivity that have retained habitat preferences of their wild conspecifics.     

Can protective attributes of artificial refuges offset predation risk in lizards?

Artificial refuges are often used to supplement habitat in areas where natural shelters have been degraded or removed. Although artificial refuges are intended to support particular species, they may be equally attractive and accessible to others, including predators. We explored the influence of snake predation risk and shelter attributes on the overnight use of different artificial refuges (timber, tiles, and iron) using the predator‐prey relationship between Boulenger's skink, Morethia boulengeri and the curl snake, Suta suta. We collected adult M. boulengeri from two bioregions in south‐eastern Australia: the Riverina, where the two species co‐occur, and the South Western Slopes, where S. suta does not occur. Two adult S. suta were collected for use as chemical donors. We conducted four experiments on overnight refuge choice to determine: (i) predator‐scent avoidance, (ii) artificial refuge preferences, (iii) a trade‐off between a preferred refuge and predator‐avoidance, and (iv) the effect of gap height on refuge preference. We found that skinks avoided predator‐scented refuges in favour of identical, but unscented refuges. Skinks preferred timber refuges, and most skinks maintained this preference when predator‐scent was added. However, when gap height was manipulated, skinks shifted to the refuge with the smallest gap. Skinks displayed complex regional variation in behaviour; skinks from both bioregions avoided predator‐scent, but in the trade‐off experiment, skinks from the South Western Slopes were less likely to avoid predator‐scented timber refuges than those from the Riverina. Our findings suggest that protective refuge attributes, such as small gap height, can offset the risk implied by predator‐scent within a refuge. This study highlights the need to consider predator‐prey interactions when designing and using artificial refuges for habitat restoration or biological monitoring.     

Sensitivity to dimethyl sulphide suggests a mechanism for olfactory navigation by seabirds

Petrels, albatrosses and other procellariiform seabirds have an excellent sense of smell, and routinely navigate over the world's oceans by mechanisms that are not well understood. These birds travel thousands of kilometres to forage on ephemeral prey patches at variable locations, yet they can quickly and efficiently find their way back to their nests on remote islands to provision chicks, even with magnetic senses experimentally disrupted. Over the seemingly featureless ocean environment, local emissions of scents released by phytoplankton reflect bathymetric features such as shelf breaks and seamounts. These features suggest an odour landscape that may provide birds with orientation cues. We have previously shown that concentrated experimental deployments of one such compound, dimethyl sulphide (DMS), attracts procellariiforms at sea, suggesting that some species can use it as a foraging cue. Here we present the first physiological demonstration that an Antarctic seabird can detect DMS at biogenic levels. We further show that birds can use DMS as an orientation cue in a non-foraging context within a concentration range that they might naturally encounter over the ocean.     

Diffusion models for population dynamics incorporating individual behavior at boundaries: applications to refuge design.

We construct models for dispersal of a population which incorporate the response of individuals to interfaces between habitat types. The models are based on random walks where there may be a bias in the direction an individual moves when it encounters an interface. This sort of dispersal process is called skew Brownian motion. Our models take the form of diffusion equations with matching conditions across the interface between regions for population densities and fluxes. We combine the dispersal models with linear population growth models which assume that the population growth rate differs between regions of different habitat types. We use those models to study issues of refuge design. We specifically consider how the effectiveness of buffer zones depends on their size, quality, and the population's response to the interface between the buffer zone and the refuge.     

Direct and indirect effects of predation on mosquitofish behavior and survival

Predation can have strong direct and indirect effects on the behavior of prey. We investigated whether predation by chain pickerel (Esox niger) caused adult eastern mosquitofish (Gambusia holbrooki) to alter their habitat use and whether pickerel predation influenced survival of adult and neonate mosquitofish. The number of adult mosquitofish using the riskier of three habitats was lowest when two predators occupied the risky habitat, intermediate in the treatment with one predator, and highest when no predators occurred there. More mosquitofish neonates survived high predation treatments than treatments lacking pickerel. We conclude that pickerel predation causes adult mosquitofish to shift to refuge habitats. The pattern of neonate survival suggests that adult habitat use may create a refuge from cannibalism for neonate mosquitofish, resulting in higher neonate survival in treatments with more pickerel. Hence, pickerel predation has a direct effect on adult mosquitofish behavior and a strong indirect effect on neonate survival. Both interspecific and intraspecific predation can effect prey populations and can interact to produce important indirect effects.     

Assessment and protective strategy of black muntjac habitat in Zhejiang, China

Habitat degradation and loss had been widely recognized as the main cause for the decline of wildlife population. Habitat assessment can provide essential information for wildlife refuge design and management. In this article, a systematic study on the black muntjac distribution, habitat quality, and spatial pattern was conducted. Initially, the main factors that influence the black muntjac habitat quality were identified, and habitat assessment criteria were set up. The vegetation, topographic, river, road, and village maps were then digitized by the geographical information system (GIS). Finally, the habitat distribution and quality were assessed based on the combination of vegetation, elevation, river system, and impact of human activities. The result showed that the potential habitat for black muntjac covered an area of 3172.6 hm2, and 501.5 hm2 of habitat in the Jiulong Mountain Natural Reserve was lost because of the degradation caused by human activities. The existing suitable habitat is 2671.1 hm2, mostly distributed in Shangliaokeng, Jiulongshan, Neibeiping, Waijiulong, and Dayanqian in Gutian, Zhejiang Province, China.     

Foraging information affects the nature of competitive interactions

If animals can learn environmental cues, how might their use of information affect competitive interactions between species? I used shared‐preference isoleg theory to generate four different predictions regarding possible ways information could affect density‐dependent habitat selection. To test these predictions, I conducted field experiments on the foraging behavior of three coexisting species of hummingbirds. I studied black‐chinned (Archilochus alexandri), blue‐throated (Lampornis clemenciae), and magnificent (Eugenes fulgens) hummingbirds at the Southwestern Research Station in the Chiricahua Mts. of Southeastern Arizona, USA. Blue‐throated hummingbirds behaviorally dominate the other two species. I found that birds foraging with complete information (via learned color cues) avoid some of the negative effects from competition. Birds that foraged with complete information remained highly selective on rich feeders even with high competitor densities. When birds suffered an information deficit, however, the two subordinate species shifted their foraging preferences. The dominant species did not shift preference. Each subordinate's shift reflects its unique place in this competitively structured guild. With high competitor densities, black‐chinneds shifted from selective to opportunistic foraging when they also suffered from an information deficit. Thus, an information deficit caused black‐chinneds to cross over their first isoleg. Rather than reduce foraging efficiency, the magnificents shortened their foraging time. I use the magnificents’ response to competition and an information deficit to explore the possibility that this guild of hummingbirds is centrifugally organized. My results require us to consider the information‐gathering (e.g., learning) abilities of individual decision‐makers when we evaluate density‐dependent habitat selection. Differences in the level of usable information can alter the way coexisting species respond to competitive interactions that potentially structure communities.     

Wall Lizards Modulate Refuge Use through Continuous Assessment of Predation Risk Level

Prey species might use several possible ways to assess predation risk when encountering a predator. Animals may consider the risk level estimated in a first encounter to remain unchanged across subsequent encounters (fixed risk response), or they may update and change their responses across encounters in accordance with short‐term changes in risk levels (flexible risk response). We examined in the field how wall lizards assess risk level by analyzing time spent in refuges after simulated predator attacks. We first examined how risk was assessed when multiple consecutive sources of risk were present simultaneously. The results suggest that wall lizards assess risk based on multiple cues, such as approach speed, directness, and persistence (measured as the distance of the predator to their refuge after an attack). When risk was high lizards remained longer in their refuges. The first decision to appear partly from the refuge depended on both approach speed and persistence, whereas the decision to emerge completely depended only on persistence and not on approach speed. This suggests that wall lizards update information on predator threat and adjusted their emergence accordingly. In a second experiment, we analyzed how short‐term changes in risk level of successive attacks affected refuge use. Successive emergence times varied as a function of current risk level of each repeated attack, independently of the risk level of previous attacks. This indicated that lizards could track short‐term changes in risk level through time and modify their initial responses when required. Fine adjustments of refuge use may help lizards to minimize costs of refuge use in unfavorable and variable environments where antipredatory responses are costly.