Risk sensitivity in three juvenile (Age-0) flatfish species: Does estuarine dependence promote risk-prone behavior?

Pleuronectid flatfish are generally thought to use stereotypical anti-predator behavior to reduce encounters with potential predators, including burial, maintaining a low profile on the bottom, cryptic coloration, and reduced activity. However, a series of laboratory experiments demonstrate significantly different predation rates on juvenile (Age-0) English sole (Parophrys vetulus), northern rock sole (Lepidopsetta polyxystra), and Pacific halibut (Hippoglossus stenolepis) by Age-2 Pacific halibut predators, suggesting differing anti-predator strategies and/or capabilities among species. In this study, behavioral attributes that control how conspicuous juvenile flatfish are to their predators, such as burial, body posture, and activity levels, were examined both in the presence and absence of perceived predation risk. English sole exhibited risk-prone behavior; they tended to bury less, exhibit an arched body posture (with head elevated off the bottom), and were more active in both the absence and presence of predators when compared to the other two species. Conversely, rock sole exhibited risk-averse behavior, being buried, inactive, with a flat body posture regardless of predation risk. Halibut demonstrated risk-sensitive behavior; behaving like English sole in the absence of predation risk, and shifted to behavior similar to that of rock sole in the face of predation risk. As an estuarine dependent species, English sole recruit to an environment that tends to be highly turbid reducing encounters with predators. As a result English sole, may have “relaxed” anti-predator behaviors in comparison to northern rock sole and Pacific halibut which recruit to less turbid coastal nurseries.     

The effect of sand and light on predation of juvenile plaice (Pleuronectes platessa) by fishes and crustaceans

Rates of predation on 0-group plaice, Pleuronectes platessa . in aquaria were compared under four different combinations of conditions to test the hypothesis that the presence of sand in which they may bury affords a refuge from predators. The effect of light and darkness on predation rate was also examined, Two crustaceans, the shrimp, Crangon crangon , and the portunid crab, Liocarcinus holatus , and two fishes, cod, Gadus morhua , and pollack, Pollachius pollachius , were used as predators. Predaton rates were significantly higher in the dark for all predators except pollack. Predation rates in the absence of sand were signifcantly greater only for pollack. The results suggest that predation rates on plaice during their juvenile nursery stage on sandy beaches will be significantly greater during darkness than during the day. Burying in sand appears to provide only a partial refuge from predation, perhaps because natural predators have evolved effective methods of foraging for buried prey.     

Adaptive flexibility in the behaviour of juvenile Atlantic salmon: short-term responses to food availability and threat from predation

Juvenile Atlantic salmon Salmo salar were shown experimentally to make adaptive behavioural decisions as a short-term response to changes in food availability and predation risk. Restricted food availability caused an increase in activity, whereas activity was decreased under predation threat. Although changes in activity were not more pronounced among the hunger-motivated fish, suggesting that they were not balancing risk and hunger, hungrier fish spent less time in refuges in the presence of a predator, indicating that they were more willing to take risks than satiated fish. Aggressive interactions among juvenile Atlantic salmon were decreased by predation threat, but were highest when predators were absent and food was abundant.     

Factors affecting the post‐release survival of cultured juvenile Pseudopleuronectes americanus

Laboratory experiments were performed with cultured and wild juvenile winter flounder Pseudopleuronectes americanus to evaluate differences in behaviour and adaptation affecting post‐release vulnerability to predation. Studies revealed that the cryptic abilities of cultured winter flounder increased over time. Sediment‐naïve, cultured fish required a minimum of 2 days to improve their burying skills and at least 90 days for colour adaptation to match the sediment. Cultured winter flounder selected sediments consisting of small grains and colours matching their own pigment. Cultured winter flounder, regardless of their colour, were significantly more vulnerable to predation by birds. Additionally, cultured fish reacted differently than wild winter flounder when exposed to cues from a potential predator.     

Sediment preferences and size-specific distribution of young-of-the-year Pacific halibut in an Alaska nursery

A combination of laboratory experiments and field surveys was used to test the hypotheses that responses to sediments change with fish size and that sediment grain-size is the predominant environmental factor affecting small-scale distribution in young-of-the-year (yoy) Pacific halibut Hippoglossus stenolepis . Laboratory tests showed that the smallest fish (31–40 mm L _T) chose fine sediments (muddy and fine sands), fish 51–70 mm had high selectivity (primarily medium sand), and the largest fish (80–150 mm) were not selective although they avoided the largest grain-sizes (pebbles and granules). Sediment preferences were correlated with size-dependent burial capabilities. Beam trawl collections were made over a 6 year period in Kachemak Bay, Alaska, to examine the distribution of yoy Pacific halibut (14–120 mm L _T) using small size classes (e.g. 10 mm intervals). Canonical correlation analysis showed that the per cent of sand in the sediment was a highly significant variable for all but one size and date combination. Catch per unit of effort (CPUE) for newly settled fish (<30 mm L _T) was highest on very fine sand, fish 41–80 mm were most abundant on fine sand, and the largest yoy fish (81–120 mm) were abundant over a range of sediments from fine sand to mud. Except for the smallest fish, Pacific halibut in the field were associated with sediments somewhat finer than predicted from the laboratory experiments; however, virtually all were captured where they could bury easily. The ability of flatfish to bury and shelter in sediment is related to fish size; consequently, habitat associations shift rapidly during the first year of life. Habitat models for yoy flatfishes should consider size-dependent shifts in capabilities and preferences.     

Sediment selection in juvenile plaice and its behavioural basis

In the laboratory juvenile plaice Pleuronectes platessa (13–114 mm) were consistently found on the finest of four sediments after 24 h both in the light and the dark. Preference when assessed both by the fish's final position and by the amount of time it spent on each sediment agreed on 72% of occasions. Size did not affect preference, which was determined principally by a fish's ability to bury in a sediment. Burial reduces activity and thereby increases the time that fish spend on sediments in which they can bury. Activity level, which is dependent on light intensity and endogenous factors, therefore determines the degree of selectivity shown. Swimming duration and resting by the largest fish (71–114 mm) did not differ among sediments. Field experiments produced results similar to those obtained in the laboratory.     

Shifting the balance between foraging and predator avoidance: the importance of food distribution for a schooling pelagic forager

It is widely held that when predator avoidance conflicts with other activities, such as feeding, avoidance of predators often takes precedence. In this study, we examine how predation risk and food distribution interact to influence the schooling behavior and swimming speed of foraging juvenile walleye pollock, Theragra chalcogramma. Fish were acclimated to either spatially and temporally clumped, or spatially and temporally dispersed food for 3 weeks. Fish were then monitored while feeding in the absence and presence of predatory sablefish, Anoplopoma fimbria. Fish foraging for clumped food swam rapidly in a loose school when predators were absent, but swam more slowly and adopted more cohesive schooling in the presence of predators, trading-off foraging opportunity for decreased vulnerability to predators. Fish foraging for dispersed food swam about slowly and did not engage in cohesive schooling in either the absence or presence of predators. These fish accepted greater predation risk in order to continue foraging, suggesting that the cost of schooling, in terms of decreased foraging opportunity, was greater when food was dispersed than when it was clumped. This lower responsiveness to predators among fish receiving dispersed food demonstrates that predator avoidance does not always take precedence over other activities, but rather, that a balance is maintained between predator avoidance and feeding, which shifts as food distribution changes.     

Experimental assessment of predation risk for juvenile green sturgeon,Acipenser medirostris,by two predatory fishes

Predation is a common cause of early life stage mortality in fishes, with reduced risk as individuals grow and become too large to be consumed by gape-limited predatory fishes. Large-bodied species, such as sturgeon, may reach this size-refuge within the first year. However, there is limited understanding of what this size threshold is despite the value of this information for conservation management. We conducted laboratory-based predation experiments on juvenile green sturgeon, Acipenser medirostris, to estimate vulnerability to predation during outmigration from their natal reaches in California to the Pacific Ocean. Two highly abundant and non-native predatory fish species (largemouth bass, Micropterus salmoides, and striped bass, Morone saxatilis) were captured in the wild to be tested with developing juvenile green sturgeon from the UC Davis Green Sturgeon Broodstock Program. Experimental tanks, each containing five predators, received thirty prey for 24-hr exposures. Between sturgeon prey trials, predators were exposed to alternative prey species to confirm predators were exhibiting normal feeding behaviors. In addition to green sturgeon mortality data, trials were video recorded and predatory behaviors were quantified. Overall, these predator species displayed much lower rates of predation on juvenile green sturgeon than alternate prey. Predation decreased with green sturgeon size, and predation risk diminished to zero once sturgeon reached a length threshold of roughly 20–22 cm total length, or between 38% and 58% of predator total length. Behavioral analyses showed low motivation to feed on green sturgeon, with both predators attempting predation less frequently as sturgeon grew. Results of this study imply that optimizing growth rates for larval and juvenile sturgeon would shorten the time in which they are vulnerable to predation. Future experiments should assess predation risk of juvenile green sturgeon by additional predator species common to the Sacramento-San Joaquin watershed.     

Predators target rare prey in coral reef fish assemblages

Predation can result in differing patterns of local prey diversity depending on whether predators are selective and, if so, how they select prey. A recent study comparing the diversity of juvenile fish assemblages among coral reefs with and without predators concluded that decreased prey diversity in the presence of predators was most likely caused by predators actively selecting rare prey species. We used several related laboratory experiments to explore this hypothesis by testing: (1) whether predators prefer particular prey species, (2) whether individual predators consistently select the same prey species, (3) whether predators target rare prey, and (4) whether rare prey are more vulnerable to predation because they differ in appearance/colouration from common prey. Rare prey suffered greater predation than expected and were not more vulnerable to predators because their appearance/colouration differed from common prey. Individual predators did not consistently select the same prey species through time, suggesting that prey selection behaviour was flexible and context dependent rather than fixed. Thus, selection of rare prey was unlikely to be explained by simple preferences for particular prey species. We hypothesize that when faced with multiple prey species predators may initially focus on rare, conspicuous species to overcome the sensory confusion experienced when attacking aggregated prey, thereby minimizing the time required to capture prey. This hypothesis represents a community-level manifestation of two well-documented and related phenomena, the “confusion effect” and the “oddity effect”, and may be an important, and often overlooked, mechanism by which predators influence local species diversity.     

The distribution of 0-group flatfish in relation to abiotic factors on the tidal flats in the brackish Dollard (Ems Estuary, Wadden Sea)

The distribution of 0-group flatfish was investigated in 1992 in the Dollard (Ems–Dollard estuary, Wadden Sea). 0-Group plaice, flounder and sole were not evenly distributed over the sampled locations. The spatial distribution pattern of 0-group flatfish in the Dollard changed during the investigation period. In the first week of sole presence, when the mean length of sole was 24–30 mm, salinity correlated significantly with sole density. The distribution of juvenile sole larger than 40 mm total length was affected by the elevation of the location: 0-group sole was restricted to the sampled site with the lowest elevation. The distribution of 0-group plaice was related to sediment: no juvenile plaice were caught at locations with more than 10% mud fraction in the sediment. The distribution of 0-group flounder was also correlated with sediment. Later in the year, salinity correlated negatively with the distribution of 0-group flounder. The influence of sediment composition is probably indirect and linked to the abundance of preferred food items, such as Corophium volutator . Abiotic conditions were suitable to 0-group plaice, flounder and sole.     

Why do juvenile fish utilise mangrove habitats?

Three hypotheses to discern the strong positive association between juvenile fish and mangrove habitat were tested with field and laboratory experiments. Artificial mangrove structure in the field attracted slightly more juvenile fish than areas without structure. Artificial structure left to accumulate fouling algae attracted four-times the total number of juvenile fish than areas without structure or areas with clean structure. Community composition of fish attracted to structure with fouling algae was different when compared with areas with no structure or clean structure; five species were attracted by structure with fouling algae whilst two species were associated with structure regardless of fouling algae. Algae were linked to increased food availability and it is suggested that this is an important selection criteria for some species. Other species were apparently attracted to structure for different reasons, and provision of shelter appears to be important. Predation pressure influenced habitat choice in small juvenile fish in laboratory experiments. In the absence of predators, small juveniles of four out of five species avoided shelter but when predators were introduced all species actively sought shelter. Large fish were apparently less vulnerable to predators and did not seek shelter when predators were added to their tank. Feeding rate was increased in the mangrove habitat for small and medium-sized fish compared with seagrass beds and mudflats indicating increased food availability or foraging efficiency within this habitat. Larger fish fed more effectively on the mudflats with an increased feeding rate in this habitat compared with adjacent habitats. The most important aspect of the mangrove habitat for small juvenile fish is the complex structure that provides maximum food availability and minimises the incidence of predation. As fish grow a shift in habitat from mangroves to mudflat is a response to changes in diet, foraging efficiency and vulnerability to predators.     

Crab shell-crushing predation and gastropod architectural defense

The shell-breaking behavior of the crabs Ozius verreauxii Saussure 1853 and Eriphia squamata, Stimpson 1859 from the Bay of Panama is described. The master claws of both these crabs are well designed for breaking shells. Small shells, relative to the size of a crab predator, are crushed by progressively breaking off larger segments of a shell's apex, while larger shells are peeled by inserting a large dactyl molar into the aperture of a shell and progressively chipping away the lip of the shell.

Heavy gastropod shells are shown to be less vulnerable to crab predators than lighter shells, and narrow shell apertures and axial shell sculpture are demonstrated to be architectural features that deter crab predation. The incidence of architectural features which deter crab predation appears to be higher for smaller gastropod species than for larger gastropods which are too large for most crab predators. Large fish predators prey upon both gastropods and shell-crushing crabs. To avoid fish predators, both these prey groups seek refuge under rocks when covered by the tide. Fish predation thus appears to enforce a close sympatry between smaller gastropods and their crab predators.     

“HKH screening”: a field bio-assessment to evaluate the ecological status of streams in the Hindu Kush-Himalayan region

Predation of fish assemblages in seagrass meadows was examined in the field and in tank experiments. Lure trolling indicated that (1) total abundance of fish was higher on bare sediment where small fish (<5 cm), including juveniles, predominated; (2) abundance was lowest in seagrass where large fish (>15 cm) predominated; (3) large ambush predators, primarily the grass goby and European eel, were almost completely restricted to seagrass; (4) the predation mode in seagrass was almost entirely ambushing or stalk-attacking, while the predation mode on bare sediment was almost entirely chase-attacking; (5) ambush predation was far more successful than chase-attack predation; and (6) overall predation risk was approximately three times higher in seagrass. Tank experiments showed that piscivory success of the grass goby was higher than that of the most common chase-attacker, the black goby, and the presence or absence of artificial seagrass, regardless of density, had no significant effect on predation success of either species. Guts of the grass goby contained food items of a wider size range that averaged twice the size of those of the black goby. Our results confirm our prediction that the risk of predation, especially of small/juvenile fish, is higher in seagrass meadows than at adjacent bare substrate, and this risk differential is explained by the presence of larger, more efficient ambush predators restricted to seagrass, and the scarcity of large chase-attack predators in the Novigrad Sea.     

Dragonfly larvae and tadpole frog space use games in varied light conditions

Predators and prey often engage in a game where predators attemptto be in areas with higher prey densities and prey attempt tobe in areas with lower predator densities. A few models havepredicted the resulting distributions of predators and prey,but little empirical data exist to test these predictions andto examine how abiotic and biotic factors shape the distributions.Thus, we observed how Anax dragonfly nymphs and Pacific treefrog tadpoles (Pseudacris regilla) either together or separatelydistributed themselves in an arena with a high- and a low-preyresource patch. Trials were conducted in high- and low-lightconditions to manipulate predation risk and to view the effectsof this abiotic factor. Counter to the model predictions, wefound that predators were not more abundant in high-resource(HR) patches, and they thus did not force prey toward beinguniformly distributed. Using a model selection approach to assesswhat factors affected predator and prey patch-switching movement,we found that prey more often left patches that had more predatorspresent, but predators surprisingly more often left patcheswith more prey present. Light levels did not affect predationrisk; however, in the dark with the associated reduction invisual information predators preferred HR patches. This causeda lower coincidence of prey and predators in patches. Predatorsalso switched patches less often when they occupied the samepatch as the other predator. This suggests that predator distributions,and indirectly prey distributions, are affected by the riskof intraguild predation.     

Mammalian predator–prey interaction in a fragmented landscape: weasels and voles

The relationship between predators and prey is thought to change due to habitat loss and fragmentation, but patterns regarding the direction of the effect are lacking. The common prediction is that specialized predators, often more dependent on a certain habitat type, should be more vulnerable to habitat loss compared to generalist predators, but actual fragmentation effects are unknown. If a predator is small and vulnerable to predation by other larger predators through intra-guild predation, habitat fragmentation will similarly affect both the prey and the small predator. In this case, the predator is predicted to behave similarly to the prey and avoid open and risky areas. We studied a specialist predator’s, the least weasel, Mustela nivalis nivalis, spacing behavior and hunting efficiency on bank voles, Myodes glareolus, in an experimentally fragmented habitat. The habitat consisted of either one large habitat patch (non-fragmented) or four small habitat patches (fragmented) with the same total area. The study was replicated in summer and autumn during a year with high avian predation risk for both voles and weasels. As predicted, weasels under radio-surveillance killed more voles in the non-fragmented habitat which also provided cover from avian predators during their prey search. However, this was only during autumn, when the killing rate was also generally high due to cold weather. The movement areas were the same for both sexes and both fragmentation treatments, but weasels of both sexes were more prone to take risks in crossing the open matrix in the fragmented treatment. Our results support the hypothesis that habitat fragmentation may increase the persistence of specialist predator and prey populations if predators are limited in the same habitat as their prey and they share the same risk from avian predation.     

Early exposure to nonlethal predation risk by size-selective predators increases somatic growth and decreases size at adulthood in three-spined sticklebacks

Predation has an important influence on life history traits in many organisms, especially when they are young. When cues of trout were present, juvenile sticklebacks grew faster. The increase in body size as a result of exposure to cues of predators was adaptive because larger individuals were more likely to survive predation. However, sticklebacks that had been exposed to cues of predators were smaller at adulthood. This result is consistent with some life history theory. However, these results prompt an alternative hypothesis, which is that the decreased size at adulthood reflects a deferred cost of early rapid growth. Compared to males, females were more likely to survive predation, but female size at adulthood was more affected by cues of predators than male size at adulthood, suggesting that size at adulthood might be more important to male fitness than to female fitness.     

Ocean acidification affects prey detection by a predatory reef fish

Changes in olfactory-mediated behaviour caused by elevated CO(2) levels in the ocean could affect recruitment to reef fish populations because larval fish become more vulnerable to predation. However, it is currently unclear how elevated CO(2) will impact the other key part of the predator-prey interaction--the predators. We investigated the effects of elevated CO(2) and reduced pH on olfactory preferences, activity levels and feeding behaviour of a common coral reef meso-predator, the brown dottyback (Pseudochromis fuscus). Predators were exposed to either current-day CO(2) levels or one of two elevated CO(2) levels (～600 μatm or ～950 μatm) that may occur by 2100 according to climate change predictions. Exposure to elevated CO(2) and reduced pH caused a shift from preference to avoidance of the smell of injured prey, with CO(2) treated predators spending approximately 20% less time in a water stream containing prey odour compared with controls. Furthermore, activity levels of fish was higher in the high CO(2) treatment and feeding activity was lower for fish in the mid CO(2) treatment; indicating that future conditions may potentially reduce the ability of the fish to respond rapidly to fluctuations in food availability. Elevated activity levels of predators in the high CO(2) treatment, however, may compensate for reduced olfactory ability, as greater movement facilitated visual detection of food. Our findings show that, at least for the species tested to date, both parties in the predator-prey relationship may be affected by ocean acidification. Although impairment of olfactory-mediated behaviour of predators might reduce the risk of predation for larval fishes, the magnitude of the observed effects of elevated CO(2) acidification appear to be more dramatic for prey compared to predators. Thus, it is unlikely that the altered behaviour of predators is sufficient to fully compensate for the effects of ocean acidification on prey mortality.     

Phenotypic variation and vulnerability to predation in juvenile bluegill sunfish (<Emphasis Type="Italic">Lepomis macrochirus</Emphasis>)

Bluegill sunfish (Lepomis macrochirus) are known to diversify into two forms specialized for foraging on either limnetic or littoral prey. Because juvenile bluegills seek vegetative cover in the presence of largemouth bass (Micropterus salmoides) predators, natural selection should favor the littoral body design at size ranges most vulnerable to predation. Yet within bluegill populations, both limnetic and littoral forms occur where vegetation and predators are present. While adaptive for foraging in different environments, does habitat-linked phenotypic variation also influence predator evasiveness for juvenile bluegills? We evaluate this question by quantifying susceptibility to predation for two groups of morphologically distinct bluegills; a limnetic form characteristic of bluegills inhabiting open water areas (limnetic bluegill) and a littoral form characteristic of bluegills inhabiting dense vegetation (littoral bluegill). In a series of predation trials, we found that bluegill behaviors differed in open water habitat but not in simulated vegetation. In open water habitat, limnetic bluegills formed more dense shoaling aggregations, maintained a larger distance from the predator, and required longer amounts of time to capture than littoral bluegill. When provided with simulated vegetation, largemouth bass spent longer amounts of time pursuing littoral bluegill and captured significantly fewer littoral bluegills than limnetic fish. Hence, morphological and behavioral variation in bluegills was linked to differential susceptibility to predation in open water and vegetated environments. Combined with previous studies, these findings show that morphological and behavioral adaptations enhance both foraging performance and predator evasiveness in different lake habitats.     

The cryptic responses of hatchery-reared sole to a natural sand substratum

In laboratory experiments, both reared and wild sole Solea solea selected a sand substratum in preference to a hard substratum. Reared sole with no previous experience of sand buried quickly when placed on sand. Light motivated burial, and the motivation of reared sole to bury was as strong as that of wild sole. The burial efficiency (the proportion of the ocular side covered by sand after a single burial attempt) of reared sole was lower than that of wild sole, but increased to that of wild sole after a period of 12 days maintenance on sand. Motivation to bury is therefore innate, but efficiency is affected by experience. The reactive distance of reared sole to a standardized predation threat was the same as that of wild sole and was shorter when buried (6 cm) than when not buried (15 cm). Burial therefore affected the response to a predation threat, indicating that burial is a cryptic behaviour. Munsell colour charts were used to determine the time required for reared sole to match the skin colour 'tone' of wild sole after placement on sand. Adaptation of colour value (lightness) took 4–7 days, but adaptation of chroma (intensity of colour) and hue took 33 and 69 days respectively. It is therefore recommended that flatfishes reared for stock enhancement exercises are conditioned to sand prior to release due to the relatively long time required for crypsis to improve through colour adaptation and burying.