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Concept: Beaver

47

Potential for habitat restoration is increasingly used as an argument for reintroducing ecosystem engineers. Beaver have well known effects on hydromorphology through dam construction, but their scope to restore wetland biodiversity in areas degraded by agriculture is largely inferred. Our study presents the first formal monitoring of a planned beaver-assisted restoration, focussing on changes in vegetation over 12years within an agriculturally-degraded fen following beaver release, based on repeated sampling of fixed plots. Effects are compared to ungrazed exclosures which allowed the wider influence of waterlogging to be separated from disturbance through tree felling and herbivory. After 12years of beaver presence mean plant species richness had increased on average by 46% per plot, whilst the cumulative number of species recorded increased on average by 148%. Heterogeneity, measured by dissimilarity of plot composition, increased on average by 71%. Plants associated with high moisture and light conditions increased significantly in coverage, whereas species indicative of high nitrogen decreased. Areas exposed to both grazing and waterlogging generally showed the most pronounced change in composition, with effects of grazing seemingly additive, but secondary, to those of waterlogging. Our study illustrates that a well-known ecosystem engineer, the beaver, can with time transform agricultural land into a comparatively species-rich and heterogeneous wetland environment, thus meeting common restoration objectives. This offers a passive but innovative solution to the problems of wetland habitat loss that complements the value of beavers for water or sediment storage and flow attenuation. The role of larger herbivores has been significantly overlooked in our understanding of freshwater ecosystem function; the use of such species may yet emerge as the missing ingredient in successful restoration.

Concepts: Biodiversity, Agriculture, Plant, Wetland, Ecosystem, Beaver, Herbivore, Biome

35

Beavers are the archetypal keystone species, which can profoundly alter ecosystem structure and function through their ecosystem engineering activity, most notably the building of dams. This can have a major impact upon water resource management, flow regimes and water quality. Previous research has predominantly focused on the activities of North American beaver (Castor canadensis) located in very different environments, to the intensive lowland agricultural landscapes of the United Kingdom and elsewhere in Europe. Two Eurasian beavers (Castor fiber) were introduced to a wooded site, situated on a first order tributary, draining from intensively managed grassland. The site was monitored to understand impacts upon water storage, flow regimes and water quality. Results indicated that beaver activity, primarily via the creation of 13 dams, has increased water storage within the site (holding ca. 1000m(3) in beaver ponds) and beavers were likely to have had a significant flow attenuation impact, as determined from peak discharges (mean 30±19% reduction), total discharges (mean 34±9% reduction) and peak rainfall to peak discharge lag times (mean 29±21% increase) during storm events. Event monitoring of water entering and leaving the site showed lower concentrations of suspended sediment, nitrogen and phosphate leaving the site (e.g. for suspended sediment; average entering site: 112±72mgl(-1), average leaving site: 39±37mgl(-1)). Combined with attenuated flows, this resulted in lower diffuse pollutant loads in water downstream. Conversely, dissolved organic carbon concentrations and loads downstream were higher. These observed changes are argued to be directly attributable to beaver activity at the site which has created a diverse wetland environment, reducing downstream hydrological connectivity. Results have important implications for beaver reintroduction programs which may provide nature based solutions to the catchment-scale water resource management issues that are faced in agricultural landscapes.

Concepts: Hydrology, Beaver, European Beaver, Beavers, Castoreum, Fur trade, North American Beaver, Keystone species

31

Globally, greenhouse gas budgets are dominated by natural sources, and aquatic ecosystems are a prominent source of methane (CH4) to the atmosphere. Beaver (Castor canadensis and Castor fiber) populations have experienced human-driven change, and CH4 emissions associated with their habitat remain uncertain. This study reports the effect of near extinction and recovery of beavers globally on aquatic CH4 emissions and habitat. Resurgence of native beaver populations and their introduction in other regions accounts for emission of 0.18-0.80 Tg CH4 year(-1) (year 2000). This flux is approximately 200 times larger than emissions from the same systems (ponds and flowing waters that became ponds) circa 1900. Beaver population recovery was estimated to have led to the creation of 9500-42 000 km(2) of ponded water, and increased riparian interface length of >200 000 km. Continued range expansion and population growth in South America and Europe could further increase CH4 emissions.

Concepts: Carbon dioxide, Natural gas, Beaver, Methane, Americas, Greenhouse gas, European Beaver, Beavers

28

Beaver have been referred to as ecosystem engineers because of the large impacts their dam building activities have on the landscape; however, the benefits they may provide to fluvial fish species has been debated. We conducted a watershed-scale experiment to test how increasing beaver dam and colony persistence in a highly degraded incised stream affects the freshwater production of steelhead (Oncorhynchus mykiss). Following the installation of beaver dam analogs (BDAs), we observed significant increases in the density, survival, and production of juvenile steelhead without impacting upstream and downstream migrations. The steelhead response occurred as the quantity and complexity of their habitat increased. This study is the first large-scale experiment to quantify the benefits of beavers and BDAs to a fish population and its habitat. Beaver mediated restoration may be a viable and efficient strategy to recover ecosystem function of previously incised streams and to increase the production of imperiled fish populations.

Concepts: River, Salmon, Oncorhynchus, Beaver, Dam, Rainbow trout, Golden trout, Brown trout

21

The Canadian beaver (Castor canadensis) is the largest indigenous rodent in North America. We report a draft annotated assembly of the beaver genome, the first for a large rodent and the first mammalian genome assembled directly from uncorrected and moderate coverage (< 30 ×) long-reads generated by single-molecule sequencing. The genome size is 2.7 Gb estimated by k-mer analysis. We assembled the beaver genome using the new Canu assembler optimized for noisy reads. The resulting assembly was refined using Pilon supported by shortreads (80 ×) and checked for accuracy by congruency against an independent short-read assembly. We scaffolded the assembly using the exon-gene models derived from 9805 full-length open reading frames (FL-ORFs) constructed from the beaver leukocyte and muscle transcriptomes. The final assembly comprised 22,515 contigs with an N50 of 278,680 bp and an N50-scaffold of 317,558 bp. Maximum contig and scaffold lengths were 3.3 and 4.2 Mb, respectively, with a combined scaffold length representing 92% of the estimated genome size. The completeness and accuracy of the scaffold assembly was demonstrated by the complete and precise exon placement for 91.1% of the 9,805 assembled FL-ORFs and 83.1% of the BUSCO (Benchmarking Universal Single-Copy Orthologs) gene set used to assess the quality of genome assemblies. Well-represented were genes involved in dentition and enamel deposition, defining characteristics of rodents with which the beaver is well-endowed. The study provides insights for genome assembly and an important genomics resource for Castoridae and rodent evolutionary biology.

Concepts: DNA, Gene, Genetics, Rodent, Beaver, Castoreum, North American Beaver, Castoridae

5

Solving problematic phylogenetic relationships often requires high quality genome data. However, for many organisms such data are still not available. Among rodents, the phylogenetic position of the beaver has always attracted special interest. The arrangement of the beaver’s masseter (jaw-closer) muscle once suggested a strong affinity to some sciurid rodents (e.g., squirrels), placing them in the Sciuromorpha suborder. Modern molecular data, however, suggested a closer relationship of beaver to the representatives of the mouse-related clade, but significant data from virtually homoplasy-free markers (for example retroposon insertions) for the exact position of the beaver have not been available. We derived a gross genome assembly from deposited genomic Illumina paired-end reads and extracted thousands of potential phylogenetically informative retroposon markers using the new bioinformatics coordinate extractor fastCOEX, enabling us to evaluate different hypotheses for the phylogenetic position of the beaver. Comparative results provided significant support for a clear relationship between beavers (Castoridae) and kangaroo rat-related species (Geomyoidea) (p < 0.0015, six markers, no conflicting data) within a significantly supported mouse-related clade (including Myodonta, Anomaluromorpha, and Castorimorpha) (p < 0.0015, six markers, no conflicting data).

Concepts: Organism, Species, Phylogenetics, Cladistics, Rodent, Beaver, Rodents, Castoridae

5

The North American Beaver (Castor canadensis) is the second largest living rodent and an iconic symbol of Canada. The beaver is a semi-aquatic browser whose diet consists of lignocellulose from a variety of plants. The beaver is a hindgut fermenter and has an enlarged ceacum that houses a complex microbiome. There have been few studies examining the microbial diversity in gastrointestinal tract of hindgut fermenting herbivores. To examine the bacterial and archaeal communities inhabiting the gastrointestinal tract of the beaver, the microbiome of the ceacum and feaces was examined using culture-independent methods. DNA from the microbial community of the ceacum and feaces of 4 adult beavers was extracted, and the16S rRNA gene was sequenced using either bacterial or archaeal specific primers. A total of 1447 and 1435 unique bacterial OTUs were sequenced from the ceacum and feaces, respectively. On average, the majority of OTUs within the ceacum were classified as Bacteroidetes (49.2%) and Firmicutes (47.6%). The feaces was also dominated by OTUs from Bacteroidetes (36.8%) and Firmicutes (58.9%). The composition of bacterial community was not significantly different among animals. The composition of the ceacal and feacal microbiome differed, but this difference is due to changes in the abundance of closely related OTUs, not because of major differences in the taxonomic composition of the communities. Within these communities, known degraders of lignocellulose were identified. In contrast, to the bacterial microbiome, the archaeal community was dominated by a single species of methanogen, Methanosphaera stadtmanae. The data presented here provide the first insight into the microbial community within the hindgut of the beaver.

Concepts: Archaea, Bacteria, Microbiology, Rodent, Beaver, Beavers, Castoreum, North American Beaver

5

Objective: The study’s purpose was to test if subclinical atherosclerosis was associated with the risk of developing HI in a large cohort of middle-aged participants. Methods: Study subjects were members of the Beaver Dam Offspring Study (BOSS), a longitudinal study of adult children of participants in the population-based Epidemiology of Hearing Loss Study (1993-present). BOSS examinations took place in 2005-2008 (baseline) and 2010-2013 (5-year follow-up). The 5-year incidence of hearing impairment was defined as a pure-tone average (PTA) of thresholds at 0.5, 1, 2 and 4 kHz > 25 dB Hearing Level (dB HL) in either ear at follow-up among participants at risk (baseline PTA in both ears < = 25 dB HL; n = 2436, mean age = 47.7 years). Atherosclerosis was measured as the mean carotid intima-media thickness and the presence of carotid artery plaque. Results: Among the 1984 participants at-risk with a follow-up audiometric examination, the 5-year incidence of hearing impairment was 8.3% (95% Confidence Interval (C.I.) 7.1, 9.5). With multivariable adjustment, carotid intima-media thickness was positively associated with hearing impairment incidence (Relative Risk (RR) = 1.14 per 0.1 mm, 95% C.I. 1.04, 1.24). The number of sites (0-6) with plaque was also positively associated with the incidence of impairment (RR = 1.16 per site, 95% C.I. 1.01, 1.32). Conclusion: Atherosclerosis was associated with the 5-year incidence of hearing impairment in this predominantly middle-aged cohort. Interventions targeting atherosclerosis prevention may help to prevent or delay the onset of hearing impairment.

Concepts: Cohort study, Longitudinal study, Epidemiology, Hearing impairment, Common carotid artery, Arithmetic mean, Intima-media thickness, Beaver

4

Through their dam-building activities and subsequent water storage, beaver have the potential to restore riparian ecosystems and offset some of the predicted effects of climate change by modulating streamflow. Thus, it is not surprising that reintroducing beaver to watersheds from which they have been extirpated is an often-used restoration and climate-adaptation strategy. Identifying sites for reintroduction, however, requires detailed information about habitat factors-information that is not often available at broad spatial scales. Here we explore the potential for beaver relocation throughout the Snohomish River Basin in Washington, USA with a model that identifies some of the basic building blocks of beaver habitat suitability and does so by relying solely on remotely sensed data. More specifically, we developed a generalized intrinsic potential model that draws on remotely sensed measures of stream gradient, stream width, and valley width to identify where beaver could become established if suitable vegetation were to be present. Thus, the model serves as a preliminary screening tool that can be applied over relatively large extents. We applied the model to 5,019 stream km and assessed the ability of the model to correctly predict beaver habitat by surveying for beavers in 352 stream reaches. To further assess the potential for relocation, we assessed land ownership, use, and land cover in the landscape surrounding stream reaches with varying levels of intrinsic potential. Model results showed that 33% of streams had moderate or high intrinsic potential for beaver habitat. We found that no site that was classified as having low intrinsic potential had any sign of beavers and that beaver were absent from nearly three quarters of potentially suitable sites, indicating that there are factors preventing the local population from occupying these areas. Of the riparian areas around streams with high intrinsic potential for beaver, 38% are on public lands and 17% are on large tracts of privately-owned timber land. Thus, although there are a large number of areas that could be suitable for relocation and restoration using beavers, current land use patterns may substantially limit feasibility in these areas.

Concepts: Hydrology, River, Stream, Riparian zone, Beaver, Land use, Geomorphology, North American Beaver

4

Establishing true phylogenetic relationships between populations is a critical consideration when sourcing individuals for translocation. This presents huge difficulties with threatened and endangered species that have become extirpated from large areas of their former range. We utilise ancient DNA (aDNA) to reconstruct the phylogenetic relationships of a keystone species which has become extinct in Britain, the Eurasian beaver Castor fiber. We sequenced seventeen 492 bp partial tRNAPro and control region sequences from Late Pleistocene and Holocene age beavers and included these in network, demographic and genealogy analyses. The mode of postglacial population expansion from refugia was investigated by employing tests of neutrality and a pairwise mismatch distribution analysis. We found evidence of a pre-Late Glacial Maximum ancestor for the Western C. fiber clade which experienced a rapid demographic expansion during the terminal Pleistocene to early Holocene period. Ancient British beavers were found to originate from the Western phylogroup but showed no phylogenetic affinity to any one modern relict population over another. Instead, we find that they formed part of a large, continuous, pan-Western European clade that harbored little internal substructure. Our study highlights the utility of aDNA in reconstructing population histories of extirpated species which has real-world implications for conservation planning.

Concepts: Biology, Species, Phylogenetic tree, Beaver, Holocene, European Beaver, Beavers, Keystone species