Calf/female ratio and population dynamics of wild forest reindeer in relation to wolf and moose abundances in a managed European ecosystem

Background The alternative prey hypothesis describes the mechanism for apparent competition whereby the mortality of the secondary prey species increases (and population size decreases decreases) by the increased predation by the shared predator if the population size of the primary prey decreases. Apparent competition is a process where the abundance of two co-existing prey species are negatively associated because they share a mutual predator, which negatively affects the abundance of both prey Here, we examined whether alternative prey and/or apparent competition hypothesis can explain the population dynamics and reproductive output of the secondary prey, wild forest reindeer (Rangifer tarandus fennicus) in Finland, in a predator-prey community in which moose (Alces alces) is the primary prey and the wolf (Canis lupus) is the generalist predator. Methods We examined a 22-year time series (1996–2017) to determine how the population size and the calf/female ratio of wild forest reindeer in Eastern Finland were related to the abundances of wolf and moose. Only moose population size was regulated by hunting. Summer predation of wolves on reindeer focuses on calves. We used least squares regression (GLS) models (for handling autocorrelated error structures and resulting pseudo-R2s) and generalized linear mixed (GLMs) models (for avoidance of negative predictions) to determine the relationships between abundances. We performed linear and general linear models for the calf/female ratio of reindeer. Results and synthesis The trends in reindeer population size and moose abundance were almost identical: an increase during the first years and then a decrease until the last years of our study period. Wolf population size in turn did not show long-term trends. Change in reindeer population size between consecutive winters was related positively to the calf/female ratio. The calf/female ratio was negatively related to wolf population size, but the reindeer population size was related to the wolf population only when moose abundance was entered as another independent variable. The wolf population was not related to moose abundance even though it is likely to consist the majority of the prey biomass. Because reindeer and moose populations were positively associated, our results seemed to support the alternative prey hypothesis more than the apparent competition hypothesis. However, these two hypotheses are not mutually exclusive and the primary mechanism is difficult to distinguish as the system is heavily managed by moose hunting. The recovery of wild forest reindeer in eastern Finland probably requires ecosystem management involving both habitat restoration and control of species abundances.

Predation of Glossiphonia weberi (Blanchard, 1897) on the invasive snail Physella acuta (Draparnaud, 1805) in the presence of an alternative prey

The North American freshwater snail Physella acuta has invaded and colonized a wide range of freshwater habitats, globally. Resembling other invasive species, P. acuta has several negative impacts on the invaded freshwater ecosystems, which calls for its regulation, preferably through biological control. The malacophagous leech Glossiphonia weberi, native to West Bengal, India, had been previously reported as a potential biocontrol agent of P. acuta. In this experiment, we observed the predation potential and prey preference of G. weberi on P. acuta in the presence of a native snail, Gyraulus convexiusculus. The results suggest that in all instances, G. weberi consumed less G. convexiusculus compared to P. acuta. In complex habitats, G. weberi showed significant prey preference to P. acuta except in the habitat characterized by the presence of pebbles. The complex habitats bear a significant (p < 0.05) negative impact only in the case of predation on P. acuta when present with conspecific but not in heterospecific prey conditions. Therefore, G. weberi qualifies as a useful biological control agent that can affect the colonization and expansion of the invasive snail P. acuta, even in the presence of alternative prey.

The dried fruit mite Carpoglyphus lactis (Acari: Carpoglyphidae) is a suitable alternative prey for Amblyseius herbicolus (Acari: Phytoseiidae)

Being generalist predators makes many members of the Phytoseiidae family a more favourable choice for biocontrol. Amblyseius herbicolus, a predator of several phytophagous pest mites and insects, has recently gained focus as an important biocontrol agent. Most studies focus on providing alternative food such as pollens as a way to increase their persistence on the targets when prey is absent. In this study, we evaluated the developmental and reproductive performance of A. herbicolus on two different diets: the dried fruit mite Carpoglyphus lactis and Raupo Typha orientalis pollen. The groups treated with C. lactis took 7 days to reach adulthood, while the groups treated with pollen took 11 days. The pollen-treated mites did not reproduce but this is likely due to the reduction of pollen quality. When given C. lactis, A. herbicolus produced more than 20 eggs over two weeks and the adults survived 18–19 days. The ratio of A. herbicolus egg to C. lactis adult used (1:20, 40 & 120) in the experiment did not affect the final population size. Cannibalism and competition seemed to affect all treatments, especially those cells started with three A. herbicolus eggs. Under laboratory conditions, A. herbicolus can be successfully cultured using C. lactis, which is an advantageous method for the laboratory and mass rearing of A. herbicolus.

Revisiting the influence of learning in predator functional response, how it can lead to shapes different from type III.

Predator/Parasitoid functional response is one of the main tools used to study predation behaviour, and in assessing the potential of biological control candidates. It is generally accepted that predator learning in prey searching and manipulation can produce the appearance of type III functional response. Holling proposed that in the presence of alternative prey, at some point the predator would shift the preferred prey, leading to the appearance of a sigmoid function that characterized that functional response. This is supported by the analogy between enzyme kinetics and functional response that Holling used as the basis for developing this theory. However, after several decades, sigmoidal functional responses appear in the absence of alternative prey in most of the biological taxa studied. Here, we propose modelling the effect of learning on the functional response by using the explicit incorporation of learning curves in the parameters of the Holling functional response, the attack rate (a), and the manipulation time (h). We then study how the variation in the parameters of the learning curves causes variations in the shape of the functional response curve. We found that the functional response product of learning can be either type I, II or III, depending on what parameters act on the organism, and how much it can learn throughout the length of the study. Therefore the presence of other types of curves should not be automatically associated with the absence of learning. These results are important from an ecological point of view because when type III functional response is associated with learning, it is generally accepted that it can operate as a stabilizing factor in population dynamics. Our results, to the contrary, suggest that depending on how it acts, it may even be destabilizing by generating the appearance of functional responses close to type I.

Resource exchange and partner recognition mediate mutualistic interactions between prey and their would-be predators

Animals may develop mutualistic associations with other species, whereby prey offer resources or services in exchange for protection from predators. Alternatively, prey may offer resources or services directly to their would-be predators in exchange for their lives. The latter may be the case of hemipterans that engage in mutualistic interactions with ants by offering a honeydew reward. We test the extent to which a honeydew offering versus partner recognition may play a role as proximate mechanisms deterring ants from predating upon their hemipteran partners. We showed that, when presented with a choice between a hemipteran partner and an alternative prey type, mutualist ants were less likely to attack and more likely to remain probing their hemipteran partners. This occurred even in the absence of an immediate sugary reward, suggesting either an evolved or learned partner recognition response. To a similar extent, however, ants were also less likely to attack the alternative prey type when laced with honey as a proxy for a honeydew reward. This was the case even after the honey had been depleted, suggesting an ability of ants to recognize new potential sources of sugars. Either possibility suggests a degree of innate or learned partner recognition.

A Comparison of Flower and Grass Strips for Augmentation of Beneficial Arthropods in Apple Orchards

Sowing plants that provide food resources in orchards is a potential habitat management practice for enhancing biological control. Flowering plants (providing pollen and nectar) and grasses (providing alternative prey) can benefit natural enemies in orchards; however, little is known about their relative importance. We studied the effect of management practices (flower strips, grass strips, and spontaneous grass) on arthropod predators under organic apple management regimes in apple orchards in Beijing, China. Orchards located at two different sites were assessed for 3 years (2017–2019). The cover crops had a significant impact on the abundance and diversity of arthropod predators. The grass treatment consistently supported significantly greater densities of alternative prey resources for predators, and predators were more abundant in the grass than in the other treatments. The Shannon–Wiener diversity was significantly higher for the cover crop treatment than for the control. Community structure was somewhat similar between the grass and control, but it differed between the flower treatment and grass/control. Weak evidence for an increase in mobile predators (ladybirds and lacewings) in the orchard canopy was found. Ladybirds and lacewings were more abundant in the grass treatment than in the other treatments in 2019 only, while the aphid abundance in the grass treatment was lowest. The fact that grass strips promoted higher predator abundance and stronger aphid suppression in comparison to the flower strips suggests that providing alternative prey for predators has great biocontrol service potential. The selection of cover crops and necessary management for conserving natural enemies in orchards are discussed in this paper.

Seasonal dynamics of a generalist and a specialist predator on a single prey

In ecological communities, the behaviour of individuals and the interaction between species may change between seasons, yet this seasonal variation is often not represented explicitly in mathematical models. As global change is predicted to alter season length and other climatic aspects, such seasonal variation needs to be included in models in order to make reasonable predictions for community dynamics. The resulting mathematical descriptions are nonautonomous models with a large number of parameters, and are therefore challenging to analyze. We present a model for two predators and one prey, whereby one predator switches hunting behaviour to seasonally include alternative prey when available. We use a combination of temporal averaging and invasion analysis to derive simplified models and determine the behaviour of the system, in particular to gain insight into conditions under which the two predators can coexist in a changing climate. We compare our results with numerical simulations of the temporally varying model.