The influence of Eurasian beaver Castor fiber activity on fish assemblages in small steppe rivers in Russia

In recent decades, the Eurasian beaver (Castor fiber) has once again become the keystone species in small river ecosystems in Russia. In many places, beaver activity has resulted in a significant change in lotic habitats, affecting the diversity, density, and biomass of aquatic organisms, including fish. While many studies have considered the ecosystem impacts of beavers, relatively few have focused on understanding the influence of beaver activity on steppe rivers. We conducted the first quantitative study of beaver impacts on fish assemblages in beaver-influenced and beaver-free sites on two small steppe rivers in the Don River basin in Russia. The presence of beavers altered the habitats in small steppe rivers and affected the diversity, density, and biomass of fish. A comparison of the number of species, density, and biomass of fish in six types of river habitats showed that these parameters were lower in beaver ponds than at riverine sites without beaver activity. Three fish species primarily preferred a single habitat type. Barbatula barbatula was found in riffles, Misgurnus fossilis in old beaver ponds, and Eudontomyzon mariae in abandoned beaver ponds. Beavers impacted fish distribution and density by changing dissolved oxygen, pH, and water current velocity. Overall, our results showed that the presence of beavers led to a temporary homogenization of fish habitats at a local scale in the valleys of small steppe rivers because beavers occupied these rivers only for a short period. However, habitat heterogeneity may increase if the beaver population stays stable or expands in the future.

     

A geospatial analysis of ecosystem engineer activity and its use during species reintroduction

The Scottish Government has announced that they are minded to allow reintroduced populations of Eurasian beaver (Castor fiber), an ecosystem engineer, to remain in Scotland. Some species and habitats of conservation importance may be affected by the activities of an ecosystem engineer. The study had two key objectives: to identify the potential location of the beaver’s two ecosystem engineering activities and to assess risks to vulnerable habitats and species. To identify potential core beaver woodland, a rule-based species distribution model was developed which included a novel method of integrating beaver territory size requirements. This was used to further predict where beavers would be unlikely to dam. We used the models to assess the spatial overlap between ecosystem engineering activities (i.e. herbivory and damming) with habitats and species of conservation importance. One hundred and five thousand five hundred eighty-six hectares of core beaver woodland was identified, which predicted the location of beaver herbivory. Our model successfully predicted the location of beaver activities surveyed in the largest beaver population in Scotland. Extensive overlap was predicted between beaver herbivory and habitats of conservation importance, in particular aspen woodland, Atlantic hazelwood, and alluvial forests. Eighty-seven percent of watercourses in Scotland were outside the beaver damming model, and hence had a lower likelihood of being dammed. The majority of freshwater pearl mussel, lamprey and Atlantic salmon Special Areas of Conservation, were in river sections less likely to be dammed. If beavers recolonise their former range in Scotland, ecosystem engineering activities are likely to be extensive. However, any impacts on vulnerable species and habitats will be in specific areas and will need to be monitored.     

Bats benefit from beavers: a facilitative link between aquatic and terrestrial food webs

Bat populations are declining in many areas, partly because up to two-thirds of their wetland habitats have been lost. One natural agent creating wetlands is the beaver, which is recolonizing its former range. Beaver flowages are known for their high production of aquatic invertebrates. We tested the hypothesis that the high numbers of insects emerging from beaver flowages influences their use by foraging bats. We compared bat use and bat numbers above flowages of introduced Canadian beavers Castor canadensis and in nearby control ponds where beavers were absent. The two bat species detected, Eptesicus nilssoni and Myotis daubentoni, used beaver flowages more than non-beaver ponds. This is especially the case for Eptesicus nilssoni. Bats also seemed to forage in larger groups while above beaver ponds compared to the control ponds. Beaver flowages appeared to improve bat habitats. A plausible reason for this could be the high number of insects emerging from beaver ponds. Favouring the beaver in habitat management is a tool for creating suitable conditions for many other species, such as bats. In areas not suited for the beaver, insect production can be achieved by imitating the beaver with man-made impoundments. This is especially important in areas which have lost most of their wetlands.     

Recovery and status of native and introduced beavers Castor fiber and Castor canadensis in France and neighbouring countries

• 1 The Eurasian beaver Castor fiber suffered a drastic reduction in both geographical range and population size, due to human persecution, until the end of the 19th century. After the adoption of protection measures, natural expansion and reintroductions led to the recovery of this species over much of its European range.
• 2 We review historical events that led to the recovery of beavers in France, and summarize the status of beavers in various river systems. Beaver establishment in France is a story of overall success: several major river systems are presently occupied, so that the species is no longer at risk in France.
• 3 However, beaver recolonization took place in parallel with increasing human impacts on the environment. In addition to natural limiting factors, anthropogenic factors impeded beaver settlement in many areas. Today, beavers often occupy suboptimal habitats and, as a consequence, come into conflict with human activities. Effective solutions for preventing beaver damage include the restoration of riparian habitats to discourage crop damage and the provision of physical barriers to protect crops.
• 4 Beaver populations reintroduced into France all originate from the relict Rhône population. However, in recent years, beavers from populations in neighbouring countries have been expanding into north‐eastern France. Therefore, our review of beaver origin and distribution in these countries may contribute to the development of appropriate national management strategies and towards important decisions, e.g. the decision to try to keep Rhône beavers genetically isolated, or to allow populations to mix.
• 5 The recently discovered presence of North American beavers Castor canadensis in three countries surrounding France has raised an important issue. This species may out‐compete C. fiber in places where the species come into contact. A programme based on field‐trapping sessions and genetic analyses has recently been initiated in some western countries in order to eradicate this non‐native species.

     

Conservation of the Eurasian beaver Castor fiber: an olfactory perspective

• 1 Chemical communication in mammals includes an array of specific behaviours that are often ignored in terms of their potential relevance to conservation. Often used during territorial or social interactions between animals, chemical communication can also be used as a tool in reintroduction programmes. Reintroductions still exhibit high failure rates and methods to improve success should be investigated. The Eurasian beaver Castor fiber has been widely reintroduced across Europe after its near extinction in the 19th century.
• 2 Using olfactory studies in the beaver, we aim to demonstrate how scent transfers a range of information about the sender which can be used to monitor social and territorial behaviour along with general well‐being. Scent manipulation can be used to reduce human–beaver conflicts, and aid reintroduction success through reducing stress and territorial conflicts, and by influencing dispersal and settlement.
• 3 Two species of beavers, the Eurasian beaver and the North American beaver Castor canadensis, occupy freshwater habitats throughout North America and in parts of South America, most of Europe and parts of Asia. Most of the reviewed literature concerns the wild Eurasian beaver, its chemical communication and conservation; however, captive studies and those addressing North American beavers are also included.
• 4 Chemical communication is advanced and has been well documented in this highly territorial species. However, few studies directly link olfaction with conservation practices.
• 5 Olfactory studies in beavers can provide non‐invasive methods to monitor translocated animals and indicators of health. We conclude that chemical analysis, olfactory studies and behavioural manipulations involving semiochemicals have important impacts on conservation and can generate practical solutions to conservation problems including aiding animal capture, captive stress reduction, breeding pair formation and release site fidelity.

     

Beavers (Castor canadensis) influence habitat for juvenile salmon in a large Alaskan river floodplain

Our aim was to determine how beavers affect habitats and food resources for juvenile salmon in the Kwethluk River in western Alaska.

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Activity of beavers as an ecological factor that affects the benthos of small rivers - a case study in the Żylica River (Poland)

The Eurasian beaver (Castor fiber) has a greater impact on local ecosystems than other herbivores, and the affected area exceeds the range of its presence. Its activity may change or create new habitats by modifying the availability of the biotic and abiotic resources available not only to beavers, but also to other groups of animals. Our aim was to study how beaver activity affects the benthos composition in a small river and in a beaver pond. The way in which beavers function in the case of small rivers has received little attention. The study showed a lower density of benthos above the dam (beaver pond - 1467 ind./m²) compared to the river (3147 ind./m²). Below the dam, the diversity of Trichoptera and Coleoptera was greater, while Diptera were more abundant in the beaver pond. Betidae were a constant component in the benthos assemblages and were most abundant in the beaver pond. Collectors-gatherers and predators were the most numerous in the beaver pond. Decreasing percentages of collectors-gatherers were observed with an increased abundance of collectors-filterers and shredders in the river. All of the ratio values except the P/T FFG (Predators to total of all other groups) were lower in the beaver pond compared to the river. The % EPT (Ephemeroptera Plecoptera and Trichoptera) was also greater in the beaver pond. Small differences in the physicochemical properties of the water and organic matter content in the bottom sediments were found both above and below the beaver dam.     

Habitat engineering by beaver benefits aquatic biodiversity and ecosystem processes in agricultural streams

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The forgotten prey of an iconic predator: a review of interactions between grey wolves Canis lupus and beavers Castor spp.

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Livestock grazing limits beaver restoration in northern New Mexico

The North American beaver (Castor canadensis) builds dams that pond water on streams, which provide crucial ecological services to aquatic and riparian ecosystems and enhance biodiversity. Consequently, there is increasing interest in restoring beavers to locations where they historically occurred, particularly in the arid western United States. However, despite often intensive efforts to reintroduce beavers into areas where they were severely reduced in numbers or eliminated due to overharvesting in the eighteenth and nineteenth centuries, beavers remain sparse or missing from many stream reaches. Reasons for this failure have not been well studied. Our goal was to evaluate certain biotic factors that may limit the occurrence of dam‐building beavers in northern New Mexico, including competitors and availability of summer and winter forage. We compared these factors at primary active dams and at control sites located in stream reaches that were physically suitable for dam‐building beavers but where none occurred. Beaver dams mostly occurred at sites that were not grazed or where there was some alternative grazing management, but were mostly absent at sites within Forest Service cattle allotments. Results indicated that cattle grazing influenced the relation between vegetation variables and beaver presence. The availability of willows (Salix spp.) was the most important plant variable for the presence of beaver dams. We conclude that grazing by cattle as currently practiced on Forest Service allotments disrupts the beaver‐willow mutualism, rendering stream reaches unsuitable for dam‐building beavers. We recommend that beaver restoration will require changes to current livestock management practices.     

Dynamics of food resources for beavers in settlements colonized and abandoned several times

The state of food resources for Eurasian beavers (Castor fiber Linnaeus, 1758) after several cycles of habitat colonization and abandonment has been analyzed in three beaver settlements on small rivers of Novgorod oblast. It has been shown that tree stands surrounding the settlements differ in composition and degree of damage but are similar in that they contain no forage for beavers: aspen is absent, and birch trees are large and require great efforts to cut them down and utilize. The undergrowth and understory also contain no aspen, while abundant willows and birch are represented by young, thin-stemmed plants unevenly distributed over the area. Tree species prevailing in abandoned settlements are not favored by beavers, implying that the animals must have expended extra energy for foraging. Supposedly, populations of beavers on small rivers in the southern taiga subzone, in habitats modified by their activities, will maintain their abundance far below the maximum possible level, depending on the rate of forage plant regrowth against the background of deteriorating habitat quality.     

The impacts of beavers Castor spp. on biodiversity and the ecological basis for their reintroduction to Scotland,UK

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The importance of aquatic vegetation in beaver diets and the seasonal and habitat specificity of aquatic‐terrestrial ecosystem linkages in a subarctic environment

The extent and ecological significance of trophic linkages across ecosystem boundaries have been the subject of considerable recent research attention. North American beavers Castor canadensis engineer terrestrial influences in aquatic ecosystems by constructing terrestrial food caches near their lodges and aquatic influences in terrestrial ecosystems by building dams and flooding low lying areas. However, it is poorly resolved to what extent beavers rely on aquatic food sources and whether this reliance is greater during winter when ice cover physically confines beavers to aquatic habitats or during summer when warm, ice free water promotes the growth and accessibility of aquatic vegetation. Working in a subarctic region, we surveyed the abundance of aquatic and terrestrial food sources in and around lotic and lentic environments and estimated their contributions to beaver diets during open water and ice covered periods using carbon and nitrogen stable isotope analysis of hair samples. Ponds had four times more aquatic vegetation than streams, but terrestrial habitats around ponds had less than half as much shrub cover as habitats adjacent to streams. Beaver diets in this subarctic environment are estimated to be comprised of 60 to 80% aquatic vegetation, with beavers occupying ponds consuming more aquatic vegetation in winter than beavers occupying streams, which rely more on terrestrial shrubs cached near their lodge. Collectively, these results show how the influence of physical barriers on ecosystem linkages can be modified by habitat‐ and season‐specific abundances of preferred resources and the potential for animals to consume food in ecosystems and seasons different from where and when the food was harvested.     

Recolonizing lost habitat—how European beavers (<Emphasis Type="Italic">Castor fiber</Emphasis>) return to south-western Germany

For the last decades, the European beaver (Castor fiber) has been recolonizing its original habitats. Reintroductions of beavers from different relict populations into southern Germany have resulted in several admixed populations, which are spreading out along various river systems. The eastern part of the German state of Baden-Württemberg is a melting pot of colonization waves originating from various introduced populations. The aim of this study was to exemplify origins and dispersal behaviour of beavers in this region using genetic fingerprint methods. Sequence analysis of hypervariable region 1 (HV1) of the mitochondrial control region and fragment length analyses at 11 microsatellite loci resulted in genetic profiles for 84 samples. The study region is being populated from three different local origins of beavers: the north of the Main-Tauber district, the Neckar River near the city of Mosbach and the Danube tributaries in the east. Main-Tauber samples were most diverse, including microsatellite alleles and HV1 haplotypes specific to C. f. albicus (from the German Elbe relict population). In view of the geographical proximity of this region to a release site of C. f. albicus in the Spessart area, this finding strongly suggests gene flow between beaver populations of different provenience. Two remaining local origins at the Neckar and Danube tributaries are closely related to each other, thus possibly descending from the same original (admixed) population. This study is intended to serve as a starting point for follow-up fine-scale research into dispersal behaviour of European beavers currently recolonizing their original habitats.     

Ecosystem engineering by invasive exotic beavers reduces in-stream diversity and enhances ecosystem function in Cape Horn,Chile

Species invasions are of global significance, but predicting their impacts can be difficult. Introduced ecosystem engineers, however, provide an opportunity to test the underlying mechanisms that may be common to all invasive engineers and link relationships between changes in diversity and ecosystem function, thereby providing explanatory power for observed ecological patterns. Here we test specific predictions for an invasive ecosystem engineer by quantifying the impacts of habitat and resource modifications caused by North American beavers (Castor canadensis) on aquatic macroinvertebrate community structure and stream ecosystem function in the Cape Horn Biosphere Reserve, Chile. We compared responses to beavers in three habitat types: (1) forested (unimpacted) stream reaches, (2) beaver ponds, and (3) sites immediately downstream of beaver dams in four streams. We found that beaver engineering in ponds created taxonomically simplified, but more productive, benthic macroinvertebrate assemblages. Specifically, macroinvertebrate richness, diversity and number of functional feeding groups were reduced by half, while abundance, biomass and secondary production increased three- to fivefold in beaver ponds compared to forested sites. Reaches downstream of beaver ponds were very similar to natural forested sections. Beaver invasion effects on both community and ecosystem parameters occurred predominantly via increased retention of fine particulate organic matter, which was associated with reduced macroinvertebrate richness and diversity (via homogenization of benthic microhabitat) and increased macroinvertebrate biomass and production (via greater food availability). Beaver modifications to macroinvertebrate community structure were largely confined to ponds, but increased benthic production in beaver-modified habitats adds to energy retention and flow for the entire stream ecosystem. Furthermore, the effects of beavers on taxa richness (negative) and measures of macroinvertebrate biomass (positive) were inversely related. Thus, while a generally positive relationship between diversity and ecosystem function has been found in a variety of systems, this work shows how they can be decoupled by responding to alterative mechanisms. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The importance of spatial scale in habitat selection by European beaver

We evaluated habitat selection by European beaver Castor fiber L. across a spatial gradient from local (within the family territory) to a broad, ecoregional scale. Based on aerial photography, we assessed the habitat composition of 150 beaver territories along the main water bodies of the Vistula River delta (northern Poland) and compared these data with 183 randomly selected sites not occupied by the species. The beavers preferred habitats with high availability of woody plants, including shrubs, and avoided anthropogenically modified habitats, such as arable lands. Within a single family territory, we observed decreasing woody plant cover with increasing distance from a colony centre, which suggests that beaver habitat preferences depend on the assessment of both the abundance and spatial distribution of preferred habitat elements. We tested the importance of spatial scale in beaver habitat selection with principal coordinates of neighbour matrices analysis, which showed that the geographical scale explained 46.7% of the variation in habitat composition, while the local beaver density explained only 10.3% of this variability. We found two main spatial gradients that were related to the broad spatial scale: first, the most important gradient was related to the largest distances between beaver sites and was independent of woody plant cover and the local beaver site density. The second most important gradient appeared more locally and was associated with these variables. Our results indicate that European beaver habitat selection was affected by different scale‐related phenomena related 1) to central place foraging behaviour, which resulted in the clumped distribution of woody plants within the territory, and 2) local population density and woody plant cover. Finally, 3) habitat selection occurs independently across the largest spatial scale studied (e.g. between watersheds), which was probably due to the limited natal dispersal range of the animals.     

Beaver invasion alters terrestrial subsidies to subantarctic stream food webs

North American beavers (Castor canadensis) were introduced to Tierra del Fuego Island in 1946 for their fur, and have since spread across the archipelago and onto the South American mainland. We assessed the impact of invasive beavers on streams of these forested watersheds by quantifying the trophic basis of production (TBP) and consumptive organic matter flows of benthic macroinvertebrate assemblages. TBP was determined in two streams: clear- and black-water. Stable isotopes were used across four streams to further elucidate food web structure and dominant pathways. TBP and stable isotopes showed that terrestrially derived organic matter (amorphous detritus, leaves, and wood) supported a majority of secondary production in the benthic food webs at all sites (forested reaches, beaver ponds, and sections downstream of ponds with foraged riparian zones). The magnitude of these flows was enhanced in beaver-modified sites compared with forested habitats (4.0–5.3× increase g AFDM m⁻² year⁻¹ in pond habitats, 1.1–2.1× increase in downstream habitats). Diatoms were the only autochthonous resource identified in macroinvertebrate guts, but their contribution to secondary production was small. Consumptive flows mirrored trends in TBP (i.e., dominance of terrestrial sources and greater magnitude in beaver ponds). Collector–gatherer consumption of amorphous detrital material dominated food web flows in all habitats, but was higher in beaver ponds relative to other habitats. Food web structure was simplified in beaver ponds; only two of the five possible functional groups contributed >1% of total organic matter flow in ponds (collector–gatherers and predators). Consumptive flows to predators increased in ponds, and stable isotopes of nitrogen and carbon (δ¹⁵N and δ¹³C) corroborated a relatively greater importance of predators (greater trophic distance), as well as less diversity of basal resources (less variation in δ¹³C) in ponds. Our findings indicate that invasive beaver’s engineering activities resulted in greater flows of terrestrial organic matter subsidies to in-stream food webs, which had a relatively greater change in the clear-water than in the black-water stream. Owing to the fact that these streams were naturally dependent on allochthonous resources for a majority of production and material flows, changes wrought by beavers to streams in forested environments are probably less than in watersheds with inherently greater dependence on autochthonous production such as the adjacent steppe biome.     

Beaver herbivory on aquatic plants

Herbivores have strong impacts on marine and terrestrial plant communities, but their impact is less well studied in benthic freshwater systems. For example, North American beavers (Castor canadensis) eat both woody and non-woody plants and focus almost exclusively on the latter in summer months, yet their impacts on non-woody plants are generally attributed to ecosystem engineering rather than herbivory. Here, we excluded beavers from areas of two beaver wetlands for over 2 years and demonstrated that beaver herbivory reduced aquatic plant biomass by 60%, plant litter by 75%, and dramatically shifted plant species composition. The perennial forb lizard’s tail (Saururus cernuus) comprised less than 5% of plant biomass in areas open to beaver grazing but greater than 50% of plant biomass in beaver exclusions. This shift was likely due to direct herbivory, as beavers preferentially consumed lizard’s tail over other plants in a field feeding assay. Beaver herbivory also reduced the abundance of the invasive aquatic plant Myriophyllum aquaticum by nearly 90%, consistent with recent evidence that native generalist herbivores provide biotic resistance against exotic plant invasions. Beaver herbivory also had indirect effects on plant interactions in this community. The palatable plant lizard’s tail was 3 times more frequent and 10 times more abundant inside woolgrass (Scirpus cyperinus) tussocks than in spatially paired locations lacking tussocks. When the protective foliage of the woolgrass was removed without exclusion cages, beavers consumed nearly half of the lizard’s tail leaves within 2 weeks. In contrast, leaf abundance increased by 73–93% in the treatments retaining woolgrass or protected by a cage. Thus, woolgrass tussocks were as effective as cages at excluding beaver foraging and provided lizard’s tail plants an associational refuge from beaver herbivory. These results suggest that beaver herbivory has strong direct and indirect impacts on populations and communities of herbaceous aquatic plants and extends the consequences of beaver activities beyond ecosystem engineering.     

A Remote Videography System for Monitoring Beavers

ABSTRACT We developed a remote videography system for monitoring behavior and demography of beavers (Castor canadensis) inside lodges and bank dens. Videography systems taped 6 beaver colonies for 1,080 hours and recorded 300 hours of beaver activity. Upon viewing videotape, we could characterize 86% of beaver activity into 1 of 12 behavioral categories. Of 26 individually marked beavers, we observed 8 (31%) on videotape and obtained a complete count of kits in 4 of 5 colonies. The remote videography system was generally useful and can provide wildlife biologists with demographic and behavioral information to support population modeling and management programs for beavers.     

Reintroducing the Eurasian beaver Castor fiber to Scotland

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