Intraguild predation and mesopredator release effect on long-lived prey

Complete extirpation of a species can generate cascading effects throughout an ecosystem, yet are precisely the goal of island eradications of pest species. “Mesopredator release effect”, an asymmetrical special case of intraguild predation, has been hypothesised as a possible indirect effect from eradications, where superpredator removal can generate a mesopredator increase which may increase the impact on their shared prey. Theoretically this suggests that for intraguild predators, the superpredator may protect the shared prey from mesopredation, and removal of superpredators alone is not recommended. We create a model of long-lived age-structured shared prey and explore the non-equilibrium dynamics of this system. The superpredator can impact all prey life-stages (adult survival and reproductive success) whereas the smaller mesopredator can only impact early life-stages (reproductive success). This model is independently tested with data from a closed oceanic island system where eradication of introduced intraguild predators is possible for conservation of threatened birds. Mesopredator release only occurs in strongly top-down moderated (resource-abundant) systems. Even when mesopredator release can occur, the negative impact of more mesopredators is outweighed by the benefit of superpredator removal, allowing recovery of the prey population. Results are robust to 10% variation in model parameters. The consideration of age-structured prey contradicts previous theoretical results for mesopredator release effect and intraguild predation. Superpredator eradication is vital for population recovery of long-lived insular species. Nonetheless island conservation must retain a whole-ecosystem perspective given the complex trophic relationships among multiple species on islands.     

Predator recognition and anti-predator responses in the rainbowfish Melanotaenia eachamensis

Predator evasion behaviour patterns of three populations of rainbowfish (Melanotaenia eachamensis) were compared. The populations differed in the level of complexity of their natural habitats and the type and extent of predation. The predator recognition abilities of fish were assessed by exposing them to models differing in their degree of predator realism. The availability of vegetated cover and the location of the models with respect to cover were manipulated. Fish from Lake Tinaroo, a relatively open habitat containing numerous predators, showed strong changes in elective group size (EGS) in response to the different models but did not rely on cover as a place of refuge. In contrast, Dirran Creek fish originate from a small, fast-flowing, structurally complex stream lacking predatory fish species, and they showed little ability to distinguish between the different models and responded to threat by spending longer in vegetated areas. Members of the Lake Eacham captive stock increased their EGS in response to models representing low threat and with more threatening models increased the amount of time spent in vegetated regions of the arena. The contrasting reactions to predatory threat displayed by these populations highlights the need to use a number of different response indices when comparing the anti-predator responses of different fish populations. These data suggest that the level of habitat complexity as well as prior predator experience influence anti-predator responses of different fish popu-lations. Received: 2 October 1996 / Accepted after revision: 27 March 1997     

Influence of prey reserve capacity on predator–prey dynamics

In this article the dynamics of a predator and prey population has been modelled when a reserve is created to protect a certain number of prey population from predation. This investigation is essential to derive some guiding principles to conserve the prey population and also to understand the behaviour and dependence of predator population on the reserve capacity. The present study concerns analysis of a fairly general model and hence some of the existing results based on specific explicit models, like Lotka–Volterra model and Rosenzweig–MacArthur model, can be derived from this work. In this study, conditions have been derived for coexistence of predator and prey, and extinction of predators. Results are obtained for global stability of required equilibrium of the model. Also, a method is suggested to compute reserve capacity in order to drive the ecosystem state to a required level. The key results developed in this article have been illustrated using numerical simulation. These results can be interpreted in different contexts like resource conservation, pest management, bio-economics of a renewable resource, etc.     

The influence of intraguild predation on prey suppression and prey release: a meta-analysis

Intraguild predation (IGP) occurs when one predator species consumes another predator species with whom it also competes for shared prey. One question of interest to ecologists is whether multiple predator species suppress prey populations more than a single predator species, and whether this result varies with the presence of IGP. We conducted a meta-analysis to examine this question, and others, regarding the effects of IGP on prey suppression. When predators can potentially consume one another (mutual IGP), prey suppression is greater in the presence of one predator species than in the presence of multiple predator species; however, this result was not found for assemblages with unidirectional or no IGP. With unidirectional IGP, intermediate predators were generally more effective than the top predator at suppressing the shared prey, in agreement with IGP theory. Adding a top predator to an assemblage generally caused prey to be released from predation, while adding an intermediate predator caused prey populations to be suppressed. However, the effects of adding a top or intermediate predator depended on the effectiveness of these predators when they were alone. Effects of IGP varied across different ecosystems (e.g., lentic, lotic, marine, terrestrial invertebrate, and terrestrial vertebrate), with the strongest patterns being driven by terrestrial invertebrates. Finally, although IGP theory is based on equilibrium conditions, data from short-term experiments can inform us about systems that are dominated by transient dynamics. Moreover, short-term experiments may be connected in some way to equilibrium models if the predator and prey densities used in experiments approximate the equilibrium densities in nature.     

Induction of anti-predator responses in the green-lipped mussel Perna viridis under hypoxia

Hypoxia due to the over enrichment of waters by nutrients is becoming a global problem. In mussels, enhanced byssus thread production is an important adaptation to the presence of crustacean predators and to energetic hydrodynamic regimes. Thread production is an energy-consuming process, so this study used the green mussel Perna viridis (L.) to examine the response to predator exposure combined with hypoxia. Hypoxia is common in sheltered bays in Hong Kong, and the mussels were collected in one such bay, Lok Wo Sha (latitude/longitude: 22o18′ N/114o10′ E) in January, 2009. The predator used in the experiments was the swimming crab Thalamita danae. Oxygen concentrations used in the 48-h experiments ranged from hypoxic to normoxic (1.5 ± 0.3 mg l⁻¹, 3.0 ± 0.3 mg l⁻¹ and 6.0 ± 0.3 mg l⁻¹). Fewer byssus threads which were also shorter and thinner were produced at reduced oxygen levels, no matter if the predator was present or not; the frequency the mussels shed stalks was also lower. Mussels exposed to the predator, however, have enhanced byssus thread production at all oxygen levels when compared with the control. This has highlighted the significance of anti-predator responses for the survival of individuals even under a stressful environment in which energy supply is limited by aerobic metabolism. Interactive effects between oxygen level and predator exposure were observed for the byssus thread production (frequency of shed stalks, mean byssus thread length, cumulative byssus thread volume), with values obtained at 1.5 and 3.0 mg O₂ l⁻¹ being statistically indistinguishable for the control group without predator but not for the predator group. The lack of differences in the byssus thread production at lower oxygen levels in the absence of predator may indicate the minimum amount of byssus that is required for settlement on a substrate.     

Footdrumming and other anti-predator responses in the bannertail kangaroo rat (Dipodomys spectabilis)

Summary Anti-predator behaviors of the bannertail kangaroo rat (Dipodomys spectabilis) toward snake predators were investigated. We induced responses by presenting a live (tethered) snake and an inflatable snake decoy to rats of known ages and sex in their territories. Comparisons of behaviors during spontaneous activity (baseline), naturally occurring snake-rat interactions, and experimental procedures further clarified anti-predator behaviors. The tethered snake immediately induced high rates of footdrumming (the repeated striking of the hind feet on the ground) in all rats (n=24) (Table 2), and individuals of both sexes and all ages footdrummed significantly more at the tethered snake than at the decoy. Other antipredator behaviors included alert postures, jumpback, kicking sand and avoidance. Juvenile rats exhibited more avoidance behavior and alert postures than adults and footdrummed at the decoy only if they experienced the snake first. We conclude that footdrumming functions as an individual alarm signal against predation by snakes in D. spectabilis. The drumming probably informs a snake that it has been detected and may cause it to leave. Because rats did not footdrum in response to aerial predators (owls), we suggest that kangaroo rats use different anti-predator strategies that depend on the kind of threat and the type of predator.     

“An eye for an eye?”—on the generality of the intimidating quality of eyespots in a butterfly and a hawkmoth

Large eyespots on the wings of butterflies and moths have been ascribed generally intimidating qualities by creating a frightening image of a bird or mammal much larger than the insect bearing the eyespots. However, evidence for this anti-predator adaptation has been largely anecdotal and only recently were peacock butterflies, Inachis io, shown to effectively thwart attacks from blue tits, Parus caeruleus. Here, we test whether large eyespots on lepidopterans are generally effective in preventing attacks from small passerines and whether the size of insect or bird can influence the outcome of interactions. We staged experiments between the larger eyed hawkmoths, Smerinthus ocellatus, and the smaller peacock butterflies, I. io, and the larger great tits, Parus major, and the smaller blue tits, P. caeruleus. Survival differed substantially between the insect species with 21 of 24 peacock butterflies, but only 6 of 27 eyed hawkmoths, surviving attacks from the birds. Thus, surprisingly, the smaller prey survived to a higher extent, suggesting that factors other than insect size may be important. However, great tits were less easily intimidated by the insects’ eyespots and deimatic behaviour and consumed 16 of 26, but the blue tits only 8 of 25, of the butterflies and hawkmoths. Our results demonstrate that eyespots per se do not guarantee survival and that these two insects bearing equally large eyespots are not equally well protected against predation.     

The effect of prey consumption on territorial defense by harriers: differential responses to neighbors versus floaters

Summary Food consumption may reduce fighting intensity of territory owners by decreasing resource value (additional food cannot be consumed) and/or increasing fighting costs (food in the digestive tract may raise injury risks). A territorial harrier's (Circus cyaneus, adult females) decision to reduce its level of aggression should depend upon whether or not the intruder was a competitor for individual prey items (as are smaller male floaters) or for the territory proper (as are female floaters and especially female neighbors). Accordingly, following meals, aggressive intensity of owners was strongly reduced towards male floaters (more were ignored), slightly reduced towards female floaters (more were called at than chased), and remained unchanged towards neighbors (virtually all were chased). Hence, alterations in aggressive behavior of territory owners following food consumption may depend upon the type of intruder and the resource under contest (a food item or a territory).     

The three-dimensional prey field of the northern krill, Meganyctiphanes norvegica, and the escape responses of their copepod prey

In the north Atlantic, Meganyctiphanes norvegica feeds predominantly on copepods, including Calanus spp. To quantify its perceptual field for prey, and the sensory systems underlying prey detection, the responses of tethered krill to free-swimming Calanus spp. were observed in 3D using silhouette video imaging. An attack–which occurred despite the krill’s being tethered—was characterized by a pronounced movement of the krill’s antennae towards the target, followed by a propulsion and opening of the feeding basket. Frequency distributions of prey detection distances were significantly different in the light vs. the dark, with median values of 26.5 mm and 19.5 mm, respectively. There were no significant differences in the angles at which prey were detected by krill (relative to the predator’s longitudinal body axis) in the light vs. the dark. Prey detections were symmetrically distributed on either side of the predator, in both light and dark. However, significant asymmetry was found in the dorsal–ventral direction with 80% of the prey detections located below the midline of the krill’s body axis and, given the placement and orientation of the compound eyes, presumably outside its visual field of view. This indicates that, at least under these conditions, vision was not the main sensory modality involved in the detection of active prey by M. norvegica. However, under some circumstances, vision may provide supplemental information. Avoidance responses of copepod prey were nearly twice the velocity of their nominal background swimming speed (153 ± 48 and 85 ± 75 mm s⁻¹, respectively), on average taking them 43 ± 16 mm away from the predator. This is far beyond the krill’s perceptual range, suggesting that the escape reaction provides an effective deterrent to predation (although perhaps less so for free-swimming krill). This information can be used to parameterize models that assess the role of krill as predators in marine ecosystems.     

The effects of opportunistic and intentional predators on the herding behavior of prey

In this article, we study how predator behavior influences the aggregation of prey into herds. Game-theoretic models of herd formation are developed based on different survival probabilities of solitary prey and prey that join the herd and on the predator's preference of what type of prey to search for. For an intentional predator that will only pursue its preferred type of prey, a single herd with no solitaries cannot emerge unless the herd acts as a prey refuge. If neither prey choice provides a refuge, it is shown that an equilibrium always exists where there are both types of prey and the predator does not always search for the same type of prey (i.e., a mixed equilibrium exists). On the other hand, if the predator is opportunistic in that it sometimes shifts to pursue the type of prey that is observed first, there may be a single herd equilibrium that does not act as a prey refuge when there is a high level of opportunistic behavior. For low opportunistic levels, a mixed equilibrium is again the only outcome. The evolutionary stability of each equilibrium is tested to see if it predicts the eventual herding behavior of prey in its corresponding model. Our analysis confirms that both predator and prey preferences (for herd or solitary) have strong effects on why prey aggregate. In particular, in our models, only the opportunistic predator can maintain all prey in a single herd that is under predation risk.     

Transient dynamics and the destabilizing effects of prey heterogeneity

The presence of prey heterogeneity and weakly interacting prey species is frequently viewed as a stabilizer of predator-prey dynamics, countering the destabilizing effects of enrichment and reducing the amplitude of population cycles. However, prior model explorations have largely focused on long-term, dynamic attractors rather than transient dynamics. Recent theoretical work shows that the presence of prey that are defended from predation can have strongly divergent effects on dynamics depending on time scale: prey heterogeneity can counteract the destabilizing effects of enrichment on predator-prey dynamics at long time scales but strongly destabilize systems during transient phases by creating long periods of low predator/prey abundance and increasing extinction probability (an effect that is amplified with increasing enrichment). We tested these general predictions using a planktonic system composed of a zooplankton predator and multiple algal prey. We first parameterized a model of our system to generate predictions and tested these experimentally. Our results qualitatively supported several model predictions. During transient phases, presence of defended algal prey increased predator extinctions at low and high enrichment levels compared to systems with only a single edible prey. This destabilizing effect was moderated at higher dilution rates, as predicted by our model. When examining dynamics beyond initial oscillations, presence of the defended prey increased predator-prey temporal variability at high nutrient enrichment but had no effect at low nutrient levels. Our results highlight the importance of considering transient dynamics when assessing the role of stabilizing factors on the dynamics of food webs.     

Evaluating the impact of caribou habitat restoration on predator and prey movement

Fragmentation of the boreal forest by linear features, including seismic lines, has destabilized predator–prey dynamics, resulting in the decline of woodland caribou (Rangifer tarandus caribou) populations. Restoration of human-altered habitat has therefore been identified as a critical management tool for achieving self-sustaining woodland caribou populations. However, only recently has testing of the response of caribou and other wildlife to restoration activities been conducted. Early work has centered around assessing changes in wildlife use of restored seismic lines. We evaluated whether restoration reduces the movement rates of predators and their associated prey, which is expected to decrease predator hunting efficiency and ultimately reduce caribou mortality. We developed a new method for using cameras to measure fine-scale movement by measuring speed as animals traveled between cameras in an array. We used our method to quantify speed of caribou, moose (Alces alces), bears (Ursus americanus), and wolves (Canis lupus) on treated (restored) and untreated seismic lines. Restoration treatments reduced travel speeds along seismic lines of wolves by 1.38 km/h, bears by 0.55 km/h, and caribou by 1.57 km/h, but did not reduce moose travel speeds. Reduced predator and caribou speeds on treated seismic lines are predicted to decrease encounter rates between predators and caribou and thus lower caribou kill rates. However, further work is needed to determine whether reduced movement rates result in reduced encounter rates with prey, and ultimately reduced caribou mortality.     

Ultrastructural observations on prey capture and digestion in the scyphomedusa Aurelia aurita

Aurelia aurita medusae are able to catch their prey with their entire body surface. Catch efficiency in medusae caught in Kiel harbour in May 1985 was found to be highest at the tentacles and lowest at the subumbrella. Surface structures of the medusa as well as the cnidom are described by SEM observations. Microbasic heterotrichous euryteles and atrichous isorhizas were found. Discharged nematocysts on the prey's skin indicate different functions of the two types. The villi in the gastral cavity show a characteristic morphological differentiation that consists of a ciliated distal and a basal area covered by vesicles. Four types of glandular cells were identified by TEM observations. Mucous cell types preferably occur in densely ciliated areas. The presence of serous cells is restricted to the basal region of the gastral villi and gastral cavity where the extracellular predigestion takes place. The time of food passage in young medusae of A. aurita decreases from 19 h at 4°C to 4 h at 22°C.     

Consequences for predators of rescue and Allee effects on prey

The size of a population can be augmented by enriching the carrying capacity of its limiting resource, or by subsidising the renewal of the resource. The well known ‘paradox of enrichment’ models the first case, in which enrichment can force consumers and their limiting resource into destabilising limit cycles, whereas impoverishment stabilises the dynamics. In this paper we model the case of resource subsidy, where the resource is a limiting prey to predators. In contrast to enrichment, the system is stabilised by an influx of prey in the form of a rescue effect, and destabilised by an outflux of prey in the form of an Allee effect. Limit cycles are not sustained by the Allee effect; instead both populations collapse to zero over a large region of the predator-prey phase plane. The catastrophic extinction of prey requires the presence of both an Allee effect on prey and a predator with a type II functional response, though neither needs to contribute a large impact to prey dynamics. The novel implication is that consumers exaggerate the impact of Allee effects on a renewing resource. Conversely, an Allee effect in the form of a cull of resource, even of small value, can trigger local extinction of resource-dependent consumers.     

Cephalopod prey of the grey-headed albatrossDiomedea chrysostoma

Cephalopod remains were collected, at regular intervals throughout the fledging period, from the stomach contents of chicks of the grey-headed albatrossDiomedea chrysostoma at Bird Island, South Georgia, in 1984 and 1986 and from regurgitations of adults at the nest in 1986. The 1984 sample was taken during a season characterised by abnormal local oceanographic conditions in which the breeding success was very low; in 1986 conditions were normal and breeding success was high. Cephalopod beaks (289 from adults; 5 651 from chicks) were identified, and allometric equations were used to estimate the biomass represented. Five cephalopod species belonging to five families (Gonatidae, Onychoteuthidae, Psychroteuthidae, Ommastrephidae and Cranchiidae) contributed 98% by number and 97% of the biomass fed to chicks. The most important species was the ommastrephidMartialia hyadesi, contributing 68.9 to 77.4% by number and 72.5 to 79.3% of the total biomass fed to chicks. The relative proportions of cephalopod species in the chicks' diet were similar between 1984 and 1986, but the total number and biomass was significantly less in 1984. There is evidence of growth ofM. hyadesi between January and June.     

Temporal variability in functional responses and prey selectivity of the pelagic mysid, Mysis mixta, in natural prey assemblages

Predation rates and prey selection of the pelagic mysid shrimp, Mysis mixta, were studied experimentally in the northern Baltic Sea in 1998 during their most intensive growth period, from June to October. Functional responses during 5 months were determined by providing the mysids with a natural zooplankton assemblage, diluted to several different concentrations. The results show that ingestion rate increased, along with mysid growth, from early summer to autumn and that saturation level was reached between 400 and 500 μg C l⁻¹. Ingestion rates increased with increasing prey concentration, and sigmoidal curves explained mostly the variation in ingestion rates (explanatory levels of 86–97%). Prey selection was evident in June, July and August, though weaker during the latter 2 months. Selection differed between the studied months but, generally, copepods were more positively selected than cladocerans. Rotifers were the main prey during June and July, when mysids were small, while larger mysids fed on copepods and cladocerans. Of the copepods, Eurytemora affinis was a truly selected species. This study shows that mysids feed on many zooplankton taxa and that they undergo ontogenetic diet shifts. Received: 19 July 2000 / Accepted: 19 October 2000     

Invasive prey impacts the abundance and distribution of native predators

While an extensive literature exists on the negative effects of invasive species, little is known about their facilitative effects on native species, particularly the role of invasives as trophic subsidies to native predators. The invasive gypsy moth (Lymantria dispar) undergoes periodic outbreaks during which it represents a super-abundant food source for predators capable of consuming it, particularly native cuckoos (Coccyzus erythropthalmus and C. americanus). We examined how gypsy moth outbreaks affect the abundance and distribution of cuckoos using the North American Breeding Bird Survey and 29 years of U.S. Forest Service gypsy moth defoliation records. Abundances of both Black-billed and Yellow-billed Cuckoos were significantly above average during outbreaks, but populations were average or below average in preceding and subsequent years, suggesting that cuckoos are immigrating to defoliations during outbreak years. Spatial analyses showed that cuckoo abundances approximately 40-150 km outside of defoliation areas were significantly below average, and these under-occupied breeding areas extended in all four compass directions around outbreaks. This result supports the idea that cuckoos locate gypsy moth outbreaks during a post-migratory nomadic phase. By shifting the annual distribution of cuckoos, gypsy moths may be shifting the trophic impact of cuckoos across large distances, which could affect native insect herbivores and plants.