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.     

Habitat selection by European beaver (Castor fiber) colonizing a boreal landscape

The beaver, exterminated in Sweden in the 19th century, was reintroduced in the 1920s. The population is still increasing its range and individuals at the expansion front can still move into optimal habitats. Therefore, there is no need for beavers to occupy marginal habitats, and preferred habitats are not yet altered by the beavers themselves. Salient features of occupied and unoccupied habitats were analysed by partial least square projections to latent structures (PLS) in order to identify the important factors. The tortuosity of streams, and occurrence of grasses and forbs in the ground layer, had the strongest positive effects. Availability of deciduous forest differed univariately between used and unused sites, but added little to the model. It is suggested that variables describing geomorphological stages in stream development will increase both the general applicability and reliability of studies that attempt to assess the quality of beaver habitat.     

A synergistic trio of invasive mammals? Facilitative interactions among beavers,muskrats, and mink at the southern end of the Americas

With ecosystems increasingly having co-occurring invasive species, it is becoming more important to understand invasive species interactions. At the southern end of the Americas, American beavers (Castor canadensis), muskrats (Ondatra zibethicus), and American mink (Neovison vison), were independently introduced. We used generalized linear models to investigate how muskrat presence related to beaver-modified habitats on Navarino Island, Chile. We also investigated the trophic interactions of the mink with muskrats and beavers by studying mink diet. Additionally, we proposed a conceptual species interaction framework involving these invasive species on the new terrestrial community. Our results indicated a positive association between muskrat presence and beaver-modified habitats. Model average coefficients indicated that muskrats preferred beaver-modified freshwater ecosystems, compared to not dammed naturally flowing streams. In addition, mammals and fish represented the main prey items for mink. Although fish were mink’s dominant prey in marine coastal habitats, muskrats represented >50 % of the biomass of mink diet in inland environments. We propose that beavers affect river flow and native vegetation, changing forests into wetlands with abundant grasses and rush vegetation. Thus, beavers facilitate the existence of muskrats, which in turn sustain inland mink populations. The latter have major impacts on the native biota, especially on native birds and small rodents. The facilitative interactions among beavers, muskrats, and mink that we explored in this study, together with other non-native species, suggest that an invasive meltdown process may exist; however further research is needed to confirm this hypothesis. Finally, we propose a community-level management to conserve the biological integrity of native ecosystems.     

Ecological engineering and aquatic connectivity: a new perspective from beaver‐modified wetlands

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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.

     

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

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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.     

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.

     

Optimal central-place foraging by beavers: Tree-size selection in relation to defensive chemicals of quaking aspen

Summary At a newly occupied pond, beavers preferentially felled aspen smaller than 7.5 cm in diameter and selected against larger size classes. After one year of cutting, 10% of the aspen had been cut and 14% of the living aspen exhibited the juvenile growth form. A phenolic compound which may act as a deterrent to beavers was found in low concentrations in aspen bark, and there was no significant regression of relative concentration of this compound on tree diameter. At a pond which had been intermittently occupied by beavers for over 20 years, beavers selected against aspen smaller than 4.5 cm in diameter, and selected in favor of aspen larger than 19.5 cm in diameter. After more than 28 years of cutting at this site, 51% of the aspen had been cut and 49% of the living aspen were juvenileform. The phenolic compound was found in significantly higher concentrations in aspen bark than at the newly occupied site, and there was a significant negative regression of relative concentration on tree diameter. The results of this study show that responses to browsing by trees place constraints on the predictive value of standard energy-based optimal foraging models, and limitations on the use of such models. Future models should attempt to account for inducible responses of plants to damage and increases in concentrations of secondary metabolites through time.     

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.     

Stichorchis subtriquetrus in European beaver from Croatia: first report

Since 1996, European beavers (Castor fiber) have been reintroduced from Germany to Croatia. However, little is known about the health status of the established population. Necropsy of a 2-year-old female European beaver, which died from a car accident, revealed 28 adult trematodes in the preserved fragments of the colon and in the peritoneal cavity. All of them were identified as Stichorchis subtriquetrus. The typical location of these parasites is the large intestine, and the finding of several specimens in the peritoneal cavity can be explained by the severe trauma. We assume that the trematode was introduced to Croatia along with the beavers. Furthermore, it is very probable that the life cycle can be completed as the intermediate hosts are part of the local fauna. This is the first record of S. subtriquetrus in Croatia.     

Abundance and habitat preferences of the southernmost population of mink: implications for managing a recent island invasion

Since 2001 invasive American mink has been known to populate Navarino Island, an island located in the pristine wilderness of the Cape Horn Biosphere Reserve, Chile, lacking native carnivorous mammals. As requested by scientists and managers, our study aims at understanding the population ecology of mink in order to respond to conservation concerns. We studied the abundance of mink in different semi-aquatic habitats using live trapping (n = 1,320 trap nights) and sign surveys (n = 68 sites). With generalized linear models we evaluated mink abundance in relation to small-scale habitat features including habitats engineered by invasive beavers (Castor canadensis). Mink have colonized the entire island and signs were found in 79% of the surveys in all types of semi-aquatic habitats. Yet, relative population abundance (0.75 mink/km of coastline) was still below densities measured in other invaded or native areas. The habitat model accuracies indicated that mink were generally less specific in habitat use, probably due to the missing limitations normally imposed by predators or competitors. The selected models predicted that mink prefer to use shrubland instead of open habitat, coastal areas with heterogeneous shores instead of flat beaches, and interestingly, that mink avoid habitats strongly modified by beavers. Our results indicate need for immediate mink control on Navarino Island. For this future management we suggest that rocky coastal shores should be considered as priority sites deserving special conservation efforts. Further research is needed with respect to the immigration of mink from adjacent islands and to examine facilitating or hampering relationships between the different invasive species present, especially if integrative management is sought.     

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.     

Concentrations of heavy metals and PCBs in the tissues of European beavers (Castor fiber) captured in northeastern Poland

European beavers (Castor fiber) from two regions were examined to identify exposure to persistent environmental contaminants. A reference group was comprised of six animals from the Forest Division of Srokowo, and an exposed group was comprised of five animals from the vicinity of a former military airport operated in 1918–1986—both from Warmia land in Poland. ¹³⁷Cs in beavers' muscles was considered negligible for the overall condition and reproduction of the animals. Elevated levels of Cd and Pb occurred in the kidneys and liver of beavers from a former military airport area. dl-PCBs were found in the liver of all specimens, and pCB no. 118 was a dominant compound. Beavers as animals living both in water and land habitats can be good bioindicators of heavy metal environmental pollution caused by human activity. Despite the significant contamination of former military sites in northeastern Poland, the beaver population shows an upward trend, reflecting the excellent adaptation of this species to a changing environment.     

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.     

Selective foraging on woody plant species by the Eurasian beaver (Castor fiber) in Telemark, Norway

Beavers Castor spp. are generalist herbivores, feeding on the bark, shoots and leaves of woody plants, terrestrial herbs and forbs, ferns and aquatic vegetation. As central-place foragers, beavers move out from water to select and cut trees and vegetation, and then transport it back to their refuge. These terrestrial forays are energetically costly; therefore, beavers should concentrate their foraging activity near the central place and increase the degree of selectivity for specific sizes or species of food with increasing distance from the water. The aim of this study was to test the predictions of the central place and the optimal foraging theories for the food selection of the Eurasian beaver Castor fiber , and show the foraging preferences of the focal species in the boreal conifer forest zone of Europe. Foraging intensity by beavers and the abundance of woody species were surveyed in transects positioned randomly at seven beaver territories. In compliance with the central-place foraging theory, the foraging intensity declined with increasing distance from the river. Beavers fed preferentially on willows ( Salix ), rowan ( Sorbus ) and birches ( Betula ), although alders ( Alnus ) dominated their diet. Size selectivity showed similar patterns to previous North American studies, which were also carried out in habitats with predominantly small saplings. The probability of selection of small saplings dropped as distance increased, which is consistent with the predictions of optimal foraging models that larger prey items are more likely to be favoured with increasing provisioning distance.     

A GIS-based habitat-suitability model as a tool for the management of beaversCastor fiber

A geographical information system (GIS)-based model, including major parameters of topography, hydrology and vegetation, was developed to assess suitable habitats for beavers in the Federal Province of Salzburg, Austria. BeaversCastor fiber Linnaeus, 1758 are on the verge of their distribution in the study area. At present, the 18 existing colonies show a rather scattered pattern, which suggests that the sub-population has not yet reached its carrying capacity, and a rapid growth phase as seen in other regions is expected. Suitable habitats have been calculated for the regions Alpenvorland, Salzburger Becken, Kalkvoralpen and Kalkhochalpen. Here optimal habitat is covering 429 ha (meeting all requirements of beavers) and suboptimal habitat is covering 187 ha (meeting all requirements except a winter food source). This represents 3.4% optimal and 1.5% sub-optimal habitat of the total available area. There are 13–14 suitable regions large enough to host at least one beaver colony. A total of 31 (23–51) colonies were calculated by the model. Based on the current sub-population of 18 colonies (36–100 animals), an average increase of 72% was expected. In an intensively managed landscape, beaver-human conflict should be anticipated and management policies established. Continuous assessment of suitable, unsuitable and sensitive habitats, extending existing conservation areas and river bank habitats, and pursuing protective measures against individuals causing conflicts are presented for the study areas.     

Beavers and lilies: selective herbivory and adaptive foraging behaviour

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Cross-species transmission of Giardia spp.: inoculation of beavers and muskrats with cysts of human, beaver, mouse, and muskrat origin

Giardia cysts isolated from humans, beavers, mice, and muskrats were tested in cross-species transmission experiments for their ability to infect either beavers or muskrats. Giardia cysts, derived from multiple symptomatic human donors and used for inoculation of beavers or muskrats, were shown to be viable by incorporation of fluorogenic dyes, excystation, and their ability to produce infections in the Mongolian gerbil model. Inoculation of beavers with 5 x 10(5) Giardia lamblia cysts resulted in the infection of 75% of the animals (n = 8), as judged by the presence of fecal cysts or intestinal trophozoites at necropsy. The mean prepatent period was 13.1 days. An infective dose experiment, using 5 x 10(1) to 5 x 10(5) viable G. lamblia cysts collected by fluorescence-activated cell sorting, demonstrated that doses of between, less than 50, and less than 500 viable cysts were required to produce infection in beavers. Scanning electron microscopy of beaver small intestine revealed that attachment of G. lamblia trophozoites produced lesions in the microvillous border. Inoculation of muskrats with G. lamblia cysts produced infections when the dose of cysts was equal to or greater than 1.25 x 10(5). The inoculation of beavers with Giardia ondatrae or Giardia muris cysts did not produce any infection; however, the administration to muskrats of Giardia cysts of beaver origin resulted in the infection of 62% of the animals (n = 8), with a prepatent period of 5 days. Our results demonstrated that beavers and muskrats could be infected with Giardia cysts derived from humans, but only by using large numbers of cysts.(ABSTRACT TRUNCATED AT 250 WORDS)     

Sympodial and monopodial branching inAcer (Aceraceae): Evolutionary trend and ecological implications

The evolutionary trend and its ecological implications in sympodial and monopodial branching patterns has been investigated in 20 JapaneseAcer spp. through comparison of shoot tip abortion and terminal bud formation. The genus is divided into two species groups according to its branching pattern, one (6 species) predominantly exhibiting sympodial branching with frequent monopodial branching in short shoots (sympodial species), and the other (14 species) exhibiting only monopodial branching (monopodial species). The early ontogeny of leaf and bud scales is described. Despite the difference in branching patterns, the bud scales of terminal buds are essentially the same in having a leaf base developed to function as a protecting organ. In all the sympodial species, during the abortion of a sympodium shoot tip, one or two pairs of primordia were found to occur on the apex, and later wither. These primordia resemble bud scales of terminal buds in their ontogeny and morphology, and appear to be rudimentary. It is suggested that a rudimentary terminal bud develops together with the establishment of sympodial branching, and that sympodial branching has originated from monopodial branching. Based on this proposed evolutionary trend, it is suggested thatAcer has moved from less shady habitats into shady habitats with monopodial branching (advantageous for vertical growth) changing into sympodial branching (advantageous for lateral spread).