High honeybee abundances reduce wild bee abundances on flowers in the city of Munich

The increase in managed honeybees (Apis mellifera) in many European cities has unknown effects on the densities of wild bees through competition. To investigate this, we monitored honeybees and non-honeybees from 01 April to 31 July 2019 and 2020 at 29 species of plants representing diverse taxonomic and floral-functional types in a large urban garden in the city of Munich in which the same plant species were cultivated in both years. No bee hives were present in the focal garden, and all bee hives in the adjacent area were closely monitored by interviewing the relevant bee keepers in both 2019 and 2020. Honeybee numbers were similar in April of both years, but increased from May to July 2020 compared to 2019. The higher densities correlated with a significant increase in shifts from wild bee to honeybee visits in May/June/July, while visitor spectra in April 2019 and 2020 remained the same. Most of the species that experienced a shift to honeybee visits in 2020 were visited mostly or exclusively for their nectar. There were no shifts towards increased wild bee visits in any species. These results from a flower-rich garden have implications for the discussion of whether urban bee keeping might negatively impact wild bees. We found clear support that high honeybee densities result in exploitative competition at numerous types of flowers.

Exploitative competition and displacement mediated by eusocial bees: experimental evidence in a wild pollinator community

Eusocial bees are likely to be ecologically important competitors for floral resources, although competitive effects can be difficult to quantify in wild pollinator communities. To investigate this, we excluded honeybees (HBE treatment), bumblebees (BBE) or both (HB&BBE) from wild-growing patches of bramble, Rubus fruticosus L. agg., flowers in two eight-day field trials at separate locations, with complementary mapping of per-site local floral resource availability. Exclusions increased per-flower volume of nectar and visitation rates of non-excluded bees, compared to control patches with no bee exclusions (CON). There was a large increase in average nectar standing crop volume both at Site 1 (+ 172%) and Site 2 (+ 137%) in HB&BBE patch flowers, and no significant change in HBE or BBE, compared to CON patches. Foraging bee responses to exclusion treatments were more pronounced at Site 2, which may be due to lower local floral resource availability, since this is likely to increase the degree of exploitative competition present. Notably, at Site 2, there was a 447% increase in larger-bodied solitary (non-Apis/Bombus) bees visiting HB&BBE patches, suggesting ecological release from competition. Hoverflies showed no response to bee removals. Numbers of other non-bee insect groups were very small and also showed no clear response to exclusions. Our findings reveal patterns of competitive exclusion between pollinator groups, mediated by resource depletion by eusocial bees. Possible long-term implications of displacement from preferred flowers, particularly where alternative forage is reduced, are discussed. Significance statement Understanding patterns of exploitative competition and displacement is necessary for pollinator conservation, particularly for vulnerable or threatened species. In this research, experimental methods reveal underlying patterns of resource competition exerted by eusocial bees in a wild pollinator community. We show that honeybees and bumblebees competitively displace each other and particularly solitary (non-Apis/Bombus) bees from bramble, an important native nectar and pollen source. Effects were stronger where local floral resource availability was identified to be limited. Notably, following experimental exclusion of both honey- and bumblebees from flowers, visitation by solitary bees increased by up to 447%, strongly suggesting ecological release from competition. These results highlight the need for informed landscape management for pollinator wellbeing, including appropriate honeybee stocking densities and improved floral resource availability.

Intraspecific competition in size-structured populations: ontogenetic shift in the importance of interference competition in a key marine herbivore

Individuals rarely have equal competitive abilities, with body size being one of the most important attributes affecting the mechanism (i.e. exploitative and interference) and consequences of competition. Competitive interactions within size-structured populations are complex and can have major implications for population dynamics, community structure and evolutionary processes. Destructive grazing of kelp beds by the green urchin Strongylocentrotus droebachiensis creates barrens where high-quality food is scarce and intraspecific competition may have an important role in structuring populations. In this study, we experimentally identified the mechanisms underlying size-asymmetric competition between small, medium, and large size classes of the green urchin. A field-based mesocosm experiment showed that small and medium sea urchins grew less and produced smaller gonads when competing for food with large conspecifics. Surprisingly, when food was provided ad libitum but large urchins were present, small individuals’ growth and foraging behavior were reduced, providing strong evidence for interference competition between small and large sea urchins. Interactions between medium and large sea urchins were, however, more influenced by exploitative competition, suggesting that sea urchins shift ontogenetically from a situation of intense interference competition to one dominated by exploitative competition. The size structure of the population can thus determine the relative importance of interference and exploitative competition. In turn, the importance of interference competition may influence size structure by inhibiting the growth of smaller urchins, a pattern consistent with the prediction of theoretical models. The consideration of size-asymmetric competitive interactions can lead to a better understanding of population size structure and dynamics.

Exploitative competition between desert kit foxes and coyotes in the Mojave Desert

Exploitative competition between two sympatric guild members can influence the composition and dynamics of an ecological community. We assessed potential exploitative competition between desert kit foxes (Vulpes macrotis arsipus) and coyotes (Canis latrans) by comparing food habits of the two species from 2009 to 2014 on a study site in the Mojave Desert in California. Desert kit foxes specialised on heteromyid rodents and invertebrates, while the most frequently occurring items consumed by coyotes were lagomorphs and rodents. Both species consumed a variety of food items throughout the study, but relative use of these items varied with year and season. Also, precipitation affected prey abundance, and this influenced prey consumption by the two species. The diets of desert kit foxes and coyotes overlapped extensively, which indicated possible exploitative competition. Desert kit foxes consistently had lower dietary diversity than coyotes, indicating that desert kit foxes are more specialist consumers while coyotes are more generalists. Dietary specialisation by kit foxes on smaller items may help reduce competition with coyotes and facilitate coexistence. Coyotes consumed anthropogenic material at much higher frequencies than desert kit foxes, and this subsidisation could increase competitive pressures on kit foxes if it results in greater coyote abundance. Desert kit foxes in California are not a protected species although formal protection has been proposed due to increasing human encroachment. Conservation strategies should include measures to maintain an abundance and diversity of natural food items and limit anthropogenic subsidisation of coyotes.

Seasonal variation in exploitative competition between honeybees and bumblebees

Honeybees (Apis mellifera) and bumblebees (Bombus spp.) often undergo exploitative competition for shared floral resources, which can alter their foraging behaviour and flower choice, even causing competitive exclusion. This may be strongest in summer, when foraging conditions are most challenging for bees, compared to other times of the year. However, the seasonal dynamics of competition between these major pollinator groups are not well understood. Here, we investigate whether the strength of exploitative competition for nectar between honeybees and bumblebees varies seasonally, and whether competitive pressure is greatest in summer months. We carried out experimental bee exclusion trials from May to late September, using experimental patches of lavender, variety Grosso, in full bloom. In each trial, we compared the numbers of honeybees (HB) foraging on patches from which bumblebees had been manually excluded (bumblebee excluded, BBE) versus control (CON) patches, HB(BBE-CON). This measure of exploitative competition varied significantly with season. As expected, mean HB(BBE-CON) was significantly greater in summer trials than in spring or autumn trials. This was despite high nectar standing crop volumes in BBE patch flowers in spring and autumn trials. Mean HB(BBE-CON) was not different between spring and autumn trials. Our results show that nectar competition between honeybees and bumblebees varies seasonally and is stronger in summer than spring or autumn, adding to current understanding of the seasonality of resource demand and competition between bee species. This information may also help to inform conservation programs aiming to increase floral resources for bees by showing when these resources are most needed.

Stuck amongst introduced species: Trophic ecology reveals complex relationships between the critically endangered Niau kingfisher and introduced predators, competitors and prey

The introduction of mammals on oceanic islands currently threatens or has caused the extinction of many endemic species. Cats and rats represent the major threat for 40 % of currently endangered island bird species. Direct (predation) and/or indirect (exploitative competition for food resource) trophic interactions are key mechanisms by which invaders cause the decrease or extinction of native populations. Here, we investigated both direct and indirect trophic interactions amongst four predator species (i.e. animals that hunt, kill and feed on other animals), including three introduced mammals (Felis silvestris catus, Rattus rattus and Rattus exulans) and one critically endangered native bird, the Niau kingfisher (Todiramphus gertrudae). All four species’ diets and prey availability were assessed from sampling at the six main kingfisher habitats on Niau Island during the breeding season. Diet analyses were conducted on 578 cat scats, 295 rat digestive tracts (218 R. exulans and 77 R. rattus) and 186 kingfisher pellets. Despite simultaneous use of morphological and PCR-based methods, no bird remains in cat and rat diet samples could be assigned to the Niau kingfisher, weakening the hypothesis of current intense predation pressure. However, we determined that Niau kingfishers mainly feed on introduced and/or cryptogenic prey and highlighted the potential for exploitative competition between this bird and both introduced rat species (for Dictyoptera, Coleoptera and Scincidae). We recommend removing the cats and both rat species, at least within kingfisher breeding and foraging areas (e.g. mechanical or chemical control, cat sterilisation, biosecurity reinforcement), to simultaneously decrease predation risk, increase key prey availability and boost kingfisher population dynamics.