Predation rates of nemertean predators: the case of a rocky shore hoplonemertean feeding on amphipods

Estimates of the predation rates of benthic nemerteans are often based on observations of single individuals, and consequently they may not be representative for all members of a population of these predators. Herein we conducted controlled and repeatable laboratory experiments on the predation rate of the hoplonemertean Amphiporus nelsoni Sánchez 1973, which is common at exposed rocky shores along the central Chilean coast. During the austral fall (April, May 2000), nemerteans were observed in relatively high numbers crawling in the intertidal zone during early morning or late-afternoon low tides. When these nemerteans were offered living amphipods held by a forceps, they immediately attacked the amphipods and fed on them. In the laboratory experiments, nemerteans preferred the amphipod Hyale maroubrae Stebbing, 1899, which is also very common in the natural habitat of A. nelsoni. The nemerteans preyed to a higher extent on small males and non-ovigerous females than would have been expected from their abundance. We suggest that these (non-reproductive) stages of H. maroubrae are very mobile and therefore have a high likelihood of encounters with nemerteans. Predation rates reached maxima when nemerteans were provided prey densities of four or more of their preferred prey species, H. maroubrae, furthermore indicating that encounter rates with prey may affect predation rates. In long-term laboratory experiments, A. nelsoni consumed more amphipods during low tide conditions than during high tide conditions. Many nemerteans in the field prefer particular environmental conditions (e.g. nocturnal low tides), which restricts the time available for successful feeding. In the long-term experiment, predation rates of A. nelsoni never exceeded 0.5 amphipods nemertean^–1 d^–1. Maximum feeding events were 3 or 4 amphipods nemertean^–1 d^–1, but this only occurred during 10 out of a possible 2634 occasions. Nemerteans that had consumed 3 or 4 amphipods during 1 day, consumed substantially less prey during the following days. Towards the end of the long-term experiment, average predation rates decreased to 0.2 amphipods nemertean^–1 d^–1, corresponding to predation rates reported for other nemertean species (0.1–0.3 prey items nemertean^–1 d^–1). We suggest that predation rates from laboratory experiments represent maximum estimates that may not be directly transferable to field populations. Additionally, low predator–prey encounter rates with preferred prey in the field may further limit the predation impact of nemertean predators in natural habitats.     

Status of the Nemertea as predators in marine ecosystems

The ecology of nemertean predators in marine ecosystems is reviewed. Nemerteans occur in most marine environments although usually in low abundances. Some species, particularly in intertidal habitats, may reach locally high densities. During specific time periods appropriate for hunting, nemerteans roam about in search of prey. Upon receiving a stimulus (usually chemical cues), many nemertean species actively pursue their prey and follow them into their dwellings or in their tracks. Other species (many hoplonemerteans) adopt a sit-and-wait strategy, awaiting prey items in strategic locations. Nemerteans possess potent neurotoxins, killing even highly mobile prey species within a few seconds and within the activity range of its attacker. Most nemertean species prey on live marine invertebrates, but some also gather on recently dead organisms to feed on them. Heteronemerteans preferentially feed on polychaetes, while most hoplonemerteans prey on small crustaceans. The species examined to date show strong preferences for selected prey species, but will attack a variety of alternative prey organisms when deprived of their favourite species. Ontogenetic changes in prey selection appear to occur, but no further information about, e.g. size selection, is available. Feeding rates as revealed from short-term laboratory experiments range on the order of 1–5 prey items d^–1. These values apparently are overestimates, since long-term experiments report substantially lower values (0.05–0.3 prey items d^–1). Nemerteans have been reported to exert a strong impact on the population size of their prey organisms through their predation activity. Considering low predation rates, these effects may primarily be a result of indirect and additive interactions. We propose future investigations on these interactive effects in combination with other predators. Another main avenue of nemertean ecological research appears to be the examination of their role in highly structured habitats such as intertidal rocky shore and coral reef environments.     

Reduction of Reactive Distance and Foraging Success in Smallmouth Bass, Micropterus dolomieu, Exposed to Elevated Turbidity Levels

We determined how turbidity affected the reactive distance and foraging success of smallmouth bass, Micropterus dolomieu. Smallmouth bass reactive distance decreased exponentially with increasing turbidity, from 65cm in clear water to 10cm at the highest turbidity. Turbidity significantly decreased the probability of a fish reacting to a prey item, but did not influence foraging success following reaction to the prey. Elevated turbidity may reduce stream fish foraging efficiency and decrease prey consumption.     

Is Foraging Time Limited During Winter? – A Feeding Experiment with Tree Sparrows Under Different Predation Risk

Small passerines are faced with a trade‐off when foraging during winter. Increasing energy reserves makes them more vulnerable to predators, while a low level of reserves exposes them to a high risk of starvation. Whether small birds under these circumstances are allowed to reduce their foraging activity under increased predation risk, for example in feeding sites more exposed to predators, remains controversial in former behavioural and ecological researches. In this study, we investigated the foraging activity of free‐living Tree Sparrow Passer montanus flocks feeding on an artificial feeding platform. The predation risk perceived by the sparrows was manipulated by placing the platform either close to or far from a bushy shelter. Foraging activity, assessed as cumulative activity of sparrows per unit time on the platform, did not differ between the low‐risk and the high‐risk conditions and did not significantly change during the day. Feeding efficiency, assessed as pecking rate, was not either reduced under the high‐risk condition. Our results suggest that sparrows were forced to feed almost continuously during the day in order to maintain their preferred level of energy reserves. However, several behavioural changes helped sparrows to adopt a safer foraging policy when feeding far from cover, as we found in another study. Altogether, sparrow flocks feeding far from cover decreased the overall foraging time (the time when any sparrow stayed on the platform) by approximately 20% as compared to the near cover condition. A possible way to maintain the same level of foraging activity despite of the reduction in overall foraging time is discussed.     

Alternative forms of competition and predation dramatically affect habitat selection under foraging--predation-risk trade-offs

Habitat selection under foraging—predation-risk trade-offshas been a frequent topic of interest to theoretical behavioraland evolutionary ecologists. However, most habitat selectionmodels assume that individuals compete exploitatively for resourcesand that predation is either density independent or dilutedcompletely by competitor number, despite empirical evidencethat other forms of competition and predation also occur innature. I developed an individual-based model for studyingthe effects of alternative forms of competition and predationon the process of habitat selection under foraging—predation-risktrade-offs. To make the model more relevant to natural populations,I assumed that individuals vary continuously in traits relatedto competitive ability and vulnerability to predation and allowedresources and predators to be distributed across more than twohabitats. The results of my investigation demonstrate thatthe predicted pattern of habitat selection can be affecteddramatically by the form predation is assumed to take. Whenpredation is density dependent or frequency dependent, individualswill tend to be distributed across habitats according to theirabsolute vulnerability to predation. In contrast, when predationis density dependent or vulnerability dependent, individualswill tend to segregate by competitive ability. Whether oneassumes that individuals compete for resources via exploitationor interference also influences the predicted pattern of habitatselection. In general, interference competition results in amore even distribution of competitors across habitats.     

The early bird gets the worm: foraging strategies of wild songbirds lead to the early discovery of food sources

Animals need to manage the combined risks of predation and starvation in order to survive. Theoretical and empirical studies have shown that individuals can reduce predation risk by delaying feeding (and hence fat storage) until late afternoon. However, little is known about how individuals manage the opposing pressures of resource uncertainty and predation risks. We suggest that individuals should follow a two-part strategy: prioritizing the discovery of food early in the day and exploiting the best patch late in the day. Using automated data loggers, we tested whether a temporal component exists in the discovery of novel foraging locations by individuals in a mixed-species foraging guild. We found that food deployed in the morning was discovered significantly more often than food deployed in the afternoon. Based on the diurnal activity patterns in this population, overall rates of new arrivals were also significantly higher than expected in the morning and significantly lower than expected in the afternoon. These results align with our predictions of a shift from patch discovery to exploitation over the course of the day.     

The response of the Eagle Owl (Bubo bubo) to an outbreak of the rabbit haemorrhagic disease

Summary The Eagle Owl (Bubo bubo) is a generalist predator that in Mediterranean areas feeds mainly on Rabbits (Oryctolagus cuniculus) At the end of 1997, a local outbreak of the rabbit haemorrhagic disease (RHD) decimated Rabbit populations in the area of Alicante (eastern Spain) so that Rabbit numbers in 1998 crashed to almost nil. Prior to the outbreak we had found 19 occupied Eagle Owl territories, and the owls had been feeding mainly on Rabbits. After the RHD epidemic, we found only six occupied territories and the owls were still feeding mainly on Rabbits. Diet composition was very similar between the periods, indicating that the owls did not diversify their diet in response to main prey scarcity. No significant differences in the proportion of Rabbit in the diet were found between the periods, suggesting that the impact of predation on Rabbits may be independent of the density of Rabbit populations (type I functional response). This response is not typically associated with generalist predators. In the territories that were still occupied after the outbreaks, the owls took substantial numbers of Hedgehogs (Erinaceus europaeus) and Red-legged Partridges (Alectoris rufa). Rats (Rattus spp), a common alternative prey, were not taken in the study area. Six breeding attempts were recorded before the RHD, and none after the RHD. This leads a to believe that only the combination of Hedgehog and Red-legged Partridge availability with the readier availability of sick Rabbits allowed a small population of owls to survive in the area, but did not allowed breeding. The predator-prey system formed by Eagle Owls and Rabbits in Mediterranean regions is more fragile than previously thought.

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Die Reaktion des Uhus (Bubo bubo) auf einen Ausbruch der hämorraghischen Krankheit bei Kaninchen (Oryctolagus cuniculus)

Zusammenfassung Der Uhu (Bubo bubo) ist ein wenig spezialisierter Greifvogel, der sich im Mittelmeerraum hauptsächlich von Kaninchen ernährt. Ende 1997 wurde die Kaninchenpopulation im Gebiet von Alicante (Ostspanien) durch einen Ausbruch der hämorrhagischen Krankheit (RHD) so stark dezimiert, dass der Kaninchenbestand 1998 nahezu zusammengebrochen war. Vor dem Ausbruch der Krankheit waren 19 Uhu-Reviere besetzt, und die Uhus lebten hauptsächlich von Kaninchen. Nach dem RHD-Ausbruch zählten wir nur noch sechs besetzte Reviere; die Uhus ernährten sich aber immer noch hauptsächlich von Kaninchen. Die Zusammensetzung der Nahrung war in beiden Perioden sehr ähnlich, was darauf hinweist, dass Uhus ihr Beutespektrum nicht erweitern, wenn ihr hauptsächliches Beutetier knapp wird. Dies legt nahe, dass die Anzahl geschlagener Kaninchen unabhängig von deren Populationsdichte ist (Typ I funktionelle Reaktion). Diese Reaktion ist untypisch für nicht spezialisierte Greifvögel. In den Revieren, die nach dem Ausbruch der Seuche noch bewohnt waren, fraßen die Uhus eine beträchtliche Anzahl Igel (Erinaceus europaeus) und Rebhühner (Alectoris rufa). Auf gewöhnliche alternative Beutetiere wie Ratten (Rattus spp) wurde im untersuchten Gebiet nicht zurückgegriffen. Sechs Brutversuche waren vor dem RHD-Ausbruch verzeichnet worden, danach kein einziger. Demzufolge lässt sich vermuten, dass nur der vereinte Effekt eines Igel- und Rebhuhnvorrats und darüber hinaus ein größeres Vorhandenseins kranker Kaninchen einer kleinen Anzahl der Uhus das Überleben in dem Gebiet ermöglichte, das Brüten jedoch nicht zuließ. Das im Mittelmeergebiet aus Uhu und Kaninchen bestehende Raubtier-Opfer-System ist anfälliger als bisher angenommen wurde.

     

Predation as a landscape effect: the trading off by prey species between predation risks and protection benefits

1. Predators impose costs on their prey but may also provide benefits such as protection against other (e.g. nest) predators. The optimal breeding location in relation to the distance from a nesting raptor varies so as to minimize the sum of costs of adult and nest predation. We provide a conceptual model to account for variation in the relative predation risks and derive qualitative predictions for how different prey species should respond to the distance from goshawk Accipiter gentilis nests. 2. We test the model predictions using a comprehensive collection of data from northern Finland and central Norway. First, we carried out a series of experiments with artificial bird nests to test if goshawks may provide protection against nest predation. Second, we conducted standard bird censuses and nest-box experiments to detect how the density or territory occupancy of several prey species varies with distance from the nearest goshawk nest. 3. Nest predation rate increased with distance from goshawk nest indicating that goshawks may provide protection for birds' nests against nest predation. Abundance (or probability of presence) of the main prey species of goshawks peaked at intermediate distances from goshawk nests, reflecting the trade-off. The abundance of small songbird species decreased with distance from goshawk nests. The goshawk poses little risk to small songbirds and they may benefit from goshawk proximity in protection against nest predation. Finally, no pattern with distance in pied flycatcher territory (nest box) occupation rate or the onset of egg-laying was detected. This is expected, as flycatchers neither suffer from marked nest predation risk nor are favoured goshawk prey. 4. Our results suggest that territory location in relation to the nest of a predator is a trade-off situation where adult birds weigh the risk of themselves being predated against the benefits accrued from increased nest survival. Prey species appear able to detect and measure alternative predation risks, and respond adaptively. From the prey perspective, the landscape is a mosaic of habitat patches the quality of which varies according to structural and floristic features, but also to the spatial distribution of predators.     

Mass regulation in response to predation risk can indicate population declines

In theory, survival rates and consequent population status might be predictable from instantaneous behavioural measures of how animals prioritize foraging vs. avoiding predation. We show, for the 30 most common small bird species ringed in the UK, that one quarter respond to higher predation risk as if it is mass-dependent and lose mass. Half respond to predation risk as if it only interrupts their foraging and gain mass thus avoiding consequent increased starvation risk from reduced foraging time. These mass responses to higher predation risk are correlated with population and conservation status both within and between species (and independently of foraging habitat, foraging guild, sociality index and size) over the last 30 years in Britain, with mass loss being associated with declining populations and mass gain with increasing populations. If individuals show an interrupted foraging response to higher predation risk, they are likely to be experiencing a high quality foraging environment that should lead to higher survival. Whereas individuals that show a mass-dependent foraging response are likely to be in lower quality foraging environments, leading to relatively lower survival.     

Optimal traits when there are several costs: the interaction of mortality and energy costs in determining foraging behavior

I analyzed the interaction of different types of costs in determiningoptimal behavior using mathematical models. The analysis concentrateson foraging behavior and asks (1) whether the cost factor thathas the greatest effect on fitness generally has the greatesteffect on optimal trait values and (2) whether increasing thesize of one type of cost makes the optimal behavior absolutelyor relatively more sensitive to that cost. The foraging costsconsidered are energy expenditure, predation risk, and othermortality factors. It is shown that increasing the magnitudeof one cost often decreases the relative and absolute sensitivityof the optimal foraging strategy to that cost. The relativefitness effects of different costs generally differ from therelative sensitivities of the optimal strategies to the costfactors. Researchers should therefore measure the shapes ofcost curves rather than their average magnitudes to determinewhich of several costs can be ignored in cost-benefit analyses.     

Behavioral interference and facilitation in the foraging cycle shape the functional response

Individual forager behaviors should affect per capita intakerates and thereby population and consumer-resource properties.We consider and incorporate conspecific facilitation and interferenceduring the separate foraging-cycle stages in a functional responsemodel that links individual behavioral interactions with consumer-resourceprocesses. Our analyses suggest that failing to properly considerand include all effects of behavioral interactions on foraging-cyclestage performances may either over- or underestimate effectsof interactions on the shape of both functional responses andpredator zero-growth isoclines. Incorporation of prey- and predator-dependentinteractions among foragers in the model produces predator isoclineswith potentials for highly complex consumer-resource dynamics.Facilitation and interference during the foraging cycle aretherefore suggested as potent behavioral mechanisms to causepatterns of community dynamics. We emphasize that correct estimationsof interaction-mediated foraging-cycle efficiencies should beconsidered in empirical and theoretical attempts to furtherour understanding of the mechanistic link between social behaviorsand higher order processes.     

The influence of potential predators on the habitat preferenda of emerging brown trout

Much research has focused on the developmental behaviour of fish and it has been shown that their sensory and physical capabilities evolve very quickly during their early life. Thus, ontogenesis could influence fishes preferences for particular environmental factors. Little is known about the habitat preferences of trout during the post-emergence phase and it is not known if they correspond to the preference curves established by the PHABSIM method for the' alevin phase'. Here, the downstream movement and habitat preferences of young emerging trout were studied in a flume. In the absence of predators, alevins preferred a water depth of 20 to 30 cm and pebble rather than gravel substratum. When emergence occurred in an area with 1⁺ trout and sculpin, Cottus gobio , almost all the emergent trout remained cryptic. When visible, most of them were in the shallowest area (10cm depth) where their preference for pebble substratum was less marked. The presence of 1⁺ trout and sculpin increased the movement downstream of young trout by 20% without changing the general and diel patterns of catches. Their presence also reduced the initial growth of 0⁺ trout.     

Angular roughshark Oxynotus centrina (Squaliformes: Oxynotidae) in captivity feeding exclusively on elasmobranch eggs: an overlooked feeding niche or a matter of individual taste?

A specimen of angular roughshark Oxynotus centrina has been kept successfully in captivity for the first time. Over a period of 24 months, the specimen preyed exclusively on the contents of elasmobranch egg cases, suggesting a specialized trophic niche.     

Size-biased predation by the gila topminnow poeciliopsis occidentalis (Baird and Girard)

The objective of this study is to document under both field and laboratory conditions the phenomenon of size-biased feeding by the Gila topminnow, Poeciliopsis occidentalis, and to shed light on the mechanisms responsible for this process. Adult female Gila topminnows feed in Monkey Spring, Santa Cruz County, Arizona, on a variety of food items, but Hyalella azteca is an important element of its invertebrate diet. The size of the amphipods ingested by female topminnows is a function of the size of the fish, with larger fish taking larger prey. There is also a strong preference for topminnows to select large amphipods from within the size limits imposed by the width of the mouth.The correlation between fish size and prey size was also seen in the laboratory, utilizing Daphnia similis and food pellets as prey. The tendency to select large prey from a range of available sizes was prominent with both Daphnia and pellets. The feeding period in the laboratory is characterized by two distinct phases. During the first phase, feeding is rapid and nonselective. As the feeding progresses, the rate of feeding drops dramatically and selectivity increases.More complex laboratory feeding experiments gave equivocal results. When fish were presented pellets of two sizes at different densities and at different ratios of small to large, size biased feeding was not clearly evident. Both predator and prey densities, agonistic behavior, and reaction distance should be explored more thoroughly as next steps in attempts to mimic feeding behavior in nature.Extracted from a Ph. D. thesis written by the junior author     

Modelling the effect of feeding-related mortality on the feeding strategy of tsetse (Diptera: Glossinidae)

Abstract. Free-living haematophagous insects risk death through host grooming responses or through increased susceptibility to predation whenever they take a bloodmeal. In this paper we investigate the effects of these risks on the feeding strategy of tsetse. A model is presented that allows for death of tsetse by starvation if they do not succeed in feeding within a fixed time (set at 6 days in the first instance) and for mortality specifically associated with feeding. In addition there is background mortality that applies to all flies at all times.
The model is used to compute the individual life-time fertility (number of female puparia per female) as a function of the probability of obtaining a meal (indicated by field data to be very high, usually > 0.85 per day) and the day on which flies start to search for a meal. We suggest that the feeding strategy that would be selected for is that which allows the maximum reproductive output. The model shows that this strategy involves making no attempts to feed for 3–4 days after the previous meal and then attempting to feed with the greatest possible probability until a meal is obtained. The predicted feeding interval, obtained independently of any trapping data, agrees closely with all previous estimates from field studies using a variety of methods. Preliminary results from a laboratory experiment reveal an increased risk of predation of recently fed as compared with hungry tsetse. The lower the actual feeding mortality the more frequently will flies be able to feed should conditions so demand. It is adaptive, however, for tsetse to delay attempting to feed for as long as they can, which is made possible by the near certainty of locating and feeding on a host within 1 day, using their sophisticated sensory systems.