Abstract Forests contribute to numerous ecosystem functions and services and contain a large proportion of terrestrial biodiversity, but they are being negatively impaced by anthropogenic activities. Forests that have never been clear‐cut and have old growth characteristics, termed “near‐natural,” often harbor different and richer species assemblages than managed forests. Alternative management strategies may be able to balance the needs of biodiversity with the demands of forestry, but evaluation efforts are limited by the challenges of measuring biodiversity. Species richness is frequently used as a simple measure of biodiversity, but research indicates that it may not adequately capture community‐level changes. Alternatively, trait‐based measures of biodiversity may prove to be useful, but research is lacking. In this paper, we use a large dataset that includes 339 obligate saproxylic beetle species collected over a decade in the boreal region throughout southern Norway to: (1) establish if there is a difference in beetle community composition between near‐natural and managed forests; and (2) determine which measures of beetle biodiversity best indicate forest naturalness. We arranged the sites in an ordination space and tested for differences in community composition between these forest types. We also tested different measures of biodiversity to determine which were the most predictive of forest naturalness. We found a clear difference in community composition between near‐natural and managed forests. Additionally, three measures of biodiversity were most predictive of forest naturalness: proportional abundance of predators, community weighted mean (CWM) of wing length, and CWM of body roundness. The probability that a forest was near‐natural increased with the proportional abundance of predators but decreased with CWM wing length and body roundness. Although species richness was higher in near‐natural forests, the effect was not significant. Overall, our findings underscore the conservation value of near‐natural forests and highlight the potential of several measures of biodiversity for determining forest quality.

Urbanization is recognized as an important driver of the diversity and abundance of tree associated insect herbivores, but its consequences for insect herbivory are controversial. A likely source of variability among studies is the insufficient consideration of intra-urban variability in forest cover. With the help of citizen scientists, we investigated the independent and interactive effect of urbanization and local canopy cover on insect herbivory in the pedunculate oak (Quercus robur) throughout most of its geographic range in Europe. The damage caused by chewing insect herbivores as well as the incidence of leaf-mining and gall-inducing herbivores consistently decreased with increasing urbanization around focal oaks. Herbivory by chewing herbivores increased with increasing forest cover, regardless of urbanization. In contrast, an increase in local canopy cover buffered the negative effect of urbanization on leaf-miners and strengthened its effect on gall-inducers. These results show the complexity of plant-herbivore interactions in urbanized areas, highlighting that the presence of local canopy cover within cities has the potential to attenuate or modify the effect of urbanization on biotic interactions.

Biodiverse communities have been shown to sustain high levels of multifunctionality and thus a loss of species likely negatively impacts ecosystem functions. For most taxa, however, roles of individual species are poorly known. Rare species, often most likely to go extinct, may have unique traits and functional roles. Alternatively, rare species may be functionally redundant, such that their loss would not disrupt ecosystem functions. We quantified the functional role of rare species by using captures of wood-living (saproxylic) beetle species, combined with recent databases of morphological and ecological traits, from three regions in central and northern Europe. Using a rarity index based on species' local abundance, geographic range, and habitat breadth, we used local and regional species removal simulations to examine the contributions of both the rarest and the most common beetle species to three measures of community functional structure: functional richness, functional specialization, and functional originality. Both regionally and locally, all three measures declined more rapidly when rare species were removed than under common (or random) species removal scenarios. These consistent patterns across scales and among forest types give evidence that rare species provide unique functional contributions, and their loss may disproportionately impact ecosystem functions. This implies that conservation measures targeting rare and endangered species, such as preserving intact forests with dead wood and mature trees, can provide broader ecosystem-level benefits. Experimental research linking functional structure to ecosystem processes should be prioritized to increase understanding of the functional consequences of species loss and to develop more effective conservation strategies.

Veteran trees are keystone structures and play vital roles in ecosystems. Tree hollows, which are filled with a mix of leaves and wood that is transformed by the biotic community into wood mould, represent an important microhabitat. Tree hollow communities consist of three major groups of organisms: microbes, mesofauna (nematodes, mites and springtails) and macrofauna (beetles, millipedes and other insects). Little is known about the interplay between associated biota and their importance for decomposition and wood mould production. The aim of this study was to test how initial food web complexity in artificial tree hollows influences community structure, decomposition, and wood mould production. We used large wooden boxes to imitate natural hollows, and mounted them on 20 veteran oaks in Southern Norway. The food webs were manipulated into three levels: (i) a complex community with macrofauna and mesofauna, (ii) a simple community without macrofauna, and (iii) only the defaunated material. Three boxes (one of each treatment, N = 60 boxes) were mounted on each oak and left in the field from May 2017 to October 2019 (28 months). After the field experiment, macrofauna and mesofauna were collected and identified, mass loss in the boxes was determined, and nitrogen and phosphorus concentrations in the wood mould were measured. We found that the complex community treatment had the greatest diversity of beetles and the highest number of predatory macrofauna. The nematodes within the complex community treatment had greater trophic structuring with K-selected species associated with stable conditions. The nematodes communities also indicated that the complex community had a balanced decomposition pathway, while the simple community was dominated by fungi and the defaunated community by bacteria. Lastly, we found that the boxes with the complex community treatment had the highest overall rate of mass loss, and greater amounts of wood mould with higher concentrations of nitrogen and phosphorus. Our results indicate that macrofauna in artificial hollows increase trophic structuring, enhance decomposition rates and wood mould production, and mediate the decomposition pathway. These findings highlight the importance of food web complexity for promoting biodiversity and ecosystem functioning in veteran tree hollows. Read the free Plain Language Summary for this article on the Journal blog.

Veteran hollow trees are keystone structures in ecosystems and provide important habitat for a diverse set of organisms, many of which are involved in the process of decomposition. Since veteran trees are ‘islands’ of high biodiversity, they provide a unique system in which to study the relationship between biodiversity and decomposition of wood. We tested this relationship with a balanced experiential design, where we quantified the taxonomic and functional diversity of beetles directly involved in the process of decomposing wood, and measured the decomposition of experimentally added bundles of small diameter wood around 20 veteran trees and 20 nearby young trees in southern Norway. We found that the diversity (both taxonomic and functional) of wood-decomposing beetles was significantly higher around the veteran trees, and beetle communities around veteran trees consisted of species with a greater preference for larger diameter wood. We extracted few beetles from the experimentally added wood bundles, regardless of the tree type that they were placed near, but decomposition rates were significantly lower around veteran trees. We speculate that slower decomposition rates around veteran trees could have been a result of a greater diversity of competing fungi, which has been found to decrease decay rates. Veteran trees provide an ecological legacy within anthropogenic landscapes, enhance biodiversity and influence wood decomposition. Actions to protect veteran trees are urgently needed in order to save these valuable organisms and their associated biodiversity.

Abstract Veteran hollow oaks (Quercus spp.) are keystone structures hosting high insect diversity but are declining in numbers due to intensification of land use and the abandonment of traditional management. The loss of this vital habitat is resulting in a reduction of biodiversity, and this likely has consequences for ecosystem functioning, especially if functional diversity is reduced. A considerable amount of research has been done on predictors of beetle taxonomic diversity in veteran oaks, but predictors of functional diversity have remained largely unexplored. The aim of this study was to establish whether the features and surroundings of veteran oaks are related to functional diversity within three functional groups of beetles (decomposers, predators, and flower visitors) and determine whether species richness and functional diversity within the groups are dependent on the same predictors. Sampling was carried out intermittently between 2004 and 2011 on 61 veteran oaks in Southern Norway. Of the 876 beetle species that were collected, 359 were determined to be decomposers, 284 were predators, and 85 were flower visitors. Species richness and functional diversity in all groups were consistently higher in traps mounted on veteran oaks in forests than in open landscapes. However, additional predictors differed between groups, and for species richness and functional diversity. Decomposer species richness responded to tree vitality, while functional diversity responded to habitat connectivity, predator species richness responded to regrowth of shrubs while functional diversity responded to tree circumference, and flower visitor richness and functional diversity did not respond to any additional predictors. Previous studies have found that the features and surroundings of veteran oaks are important for conservation of taxonomic diversity, and the results from this study indicate that they are also important for functional diversity within multiple functional groups.

Predation of invertebrate pest by natural enemies is a critical contribution of nature to people, because invertebrate pests cause a vast amount of economic damage and pesticides use has many long-term costs. Veteran trees are keystone structures and hotspots for biodiversity, and are a potential source of natural enemies. To explore this, we used a balanced experimental design where we measured predatory beetle diversity and attack marks on three colors of artificial caterpillars placed around 20 veteran oaks and 20 nearby young oaks, in Southern Norway. We predicted that around the veteran oaks there would be a greater diversity of predatory beetles and more invertebrate attacks on artificial caterpillars. Sampling for predatory beetles was conducted in summer 2017 and 2018, and invertebrate attacks were measured in 2018. We found support for the predictions: diversity of predatory beetles was higher around veteran trees and there were more arthropod attack marks on artificial caterpillars placed around veteran trees. Our results indicated that veteran trees are a source of natural enemies. Valuing and protecting veteran trees and their communities is an essential step towards a more sustainable system of management that has the possibility of promoting both the wellbeing of people and biodiversity.

Parasites have detrimental effects on their hosts’ fitness. Therefore, behavioural adaptations have evolved to avoid parasites or, when an individual is already in contact with a parasite, prevent or minimize infections. Such anti‐parasite behaviours can be very effective, but can also be costly for the host. Specifically, ectoparasites can elicit strong host anti‐parasite behaviours and interactions between fleas (Siphonaptera) and their hosts are one of the best studied. In altricial bird species, nest fleas can negatively affect both parent and offspring fitness components. However, knowledge on the effects of fleas on precocial bird species is scarce. Research on geese in the Canadian Arctic indicated that fleas have a negative impact on reproductive success. One possible hypothesis is that fleas may affect female incubation behaviour. Breeding females with many fleas in their nest may increase the frequency and/or duration of incubation breaks and could even totally desert their nest. The aim of our study was to 1) determine if a similar negative relationship existed between flea abundance and reproductive success in our study colony of Arctic breeding barnacle geese Branta leucopsis and 2) experimentally quantify if such effects could be explained by a negative effect of nest fleas on female behaviour. We compared host anti‐parasite and incubation behaviour between experimentally flea‐reduced and control nests using wildlife cameras and temperature loggers. We found that flea abundance was negatively associated with hatching success. We found little experimental support, however, for changes in behaviour of the breeding female as a possible mechanism to explain this effect.

Arctic terrestrial ecosystems are ideal systems to study host-parasite interactions because they are sensitive and have fewer confounding interactions than lower latitudes. Since the Arctic is experiencing rapid climate change, research there can provide models to better understand and predict future changes in other systems. The Barnacle goose (Branta leucopsis) and its ectoparasite the flea (Ceratophyllus vagabundus vagabundus) is an ideal Arctic host-parasite system to study because Barnacle geese have important roles in Arctic ecosystems as selective grazers and prey items and have been intensively studied for decades. Also, recent studies suggest that Arctic avian flea infestations may be increasing, possibly due to climate change. However, few studies have considered the effect of fleas on Arctic nesting birds and little is known of Arctic avian flea ecology, most importantly whether they overwinter in the High Arctic or are reintroduced each summer by migrating birds. Furthermore, sampling methods for avian fleas are time-consuming and semi-invasive to their hosts. Therefore, the project had four aims: (i) contribute to the knowledge of Arctic flea ecology by looking for evidence of overwintering, (ii) use an experimental study to assess the impact of flea infestations on the hatching success of Barnacle goose eggs, (iii) assess the simple and disturbance-free method of estimating fleas with photographs of blood spots on goose eggs, (iv) and test the results of both the experimental study as well as the application of blood spots as a proxy measure for flea infestations through a larger observational study. It was found that fleas were likely overwintering in empty nest bowls and some evidence indicated that they might be capable of a two-year life cycle in the High Arctic. The findings from the experimental study indicated that heavy flea infestations had a negative impact on the hatching success of Barnacle goose eggs. It was also determined that blood spots were a good proxy measure of flea infestations, but become less accurate through incubation due to wear from incubating birds. Finally, the observational study supported the findings of the experimental study and demonstrated the power and simplicity of using blood spots as a proxy measure for flea infestations. Based on these findings, it is suggested that researchers interested in either reproductive success or incubation behavior of Barnacle geese should include a measure of flea infestations in their studies and using blood spots as a proxy measure is a simple and disturbance-free method of doing this.