Composition Vendor Newgen KnowledgeWorks (P) Ltd. PLoS One plos plosone PLOS One 1932-6203 Public Library of Science San Francisco, CA USA 10.1371/journal.pone.0349313 PONE-D-25-54787 Research Article Biology and life sciencesOrganismsEukaryotaAnimalsVertebratesAmniotesMammalsRodentsBeavers Biology and life sciencesZoologyAnimalsVertebratesAmniotesMammalsRodentsBeavers Ecology and environmental sciencesAquatic environmentsFreshwater environmentsMarshes Earth sciencesMarine and aquatic sciencesAquatic environmentsFreshwater environmentsMarshes Earth sciencesGeophysicsTides Physical sciencesPhysicsGeophysicsTides Biology and life sciencesEcologyEcosystemsWetland ecosystemsDelta ecosystems Ecology and environmental sciencesEcologyEcosystemsWetland ecosystemsDelta ecosystems Ecology and environmental sciencesAquatic environmentsFreshwater environmentsFresh water Earth sciencesMarine and aquatic sciencesAquatic environmentsFreshwater environmentsFresh water Ecology and environmental sciencesAquatic environmentsFreshwater environmentsWetlands Earth sciencesMarine and aquatic sciencesAquatic environmentsFreshwater environmentsWetlands Earth sciencesGeomorphologyTopographyLandformsWetlands Earth sciencesMarine and aquatic sciencesBodies of waterRivers Ecology and environmental sciencesAquatic environmentsFreshwater environmentsRivers Earth sciencesMarine and aquatic sciencesAquatic environmentsFreshwater environmentsRivers Earth sciencesHydrologyFlooding Beaver in tidal habitat: Examples from the Pacific Northwest Beaver in tidal habitat https://orcid.org/0000-0001-6286-7935 Hood W. Gregory Conceptualization Data curation Formal analysis Funding acquisition Investigation Methodology Project administration Writing – original draft Writing – review & editing * Skagit River System Cooperative, Burlington, Washington, United States of America Laruelle Goulven G Editor Universite Libre de Bruxelles, BELGIUM

The author has declared that no competing interests exist.

 * E-mail: ghood@skagitcoop.org 872026 2026 21 7 e0349313 9102025 2842026 2026 W. Gregory Hood This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Beaver are typically considered fluvial or lacustrine animals, often converting lotic habitats into lentic ones with their dams. This results in extensive changes in ecosystem structure and processes so that beaver are considered the quintessential ecosystem engineer. Here I broaden our appreciation of the adaptability of beaver by describing their widespread presence in tidal river deltas and estuaries of the Pacific Northwest (coastal British Columbia, Washington, and Oregon), where tides can range between 1.5 and 5.0 m. These observations expand the known habitat distribution of beaver and invite investigation of the ecosystem consequences of beaver in tidal wetlands. In these oligohaline to fresh tidal systems, channel profile surveys with real-time kinematic (RTK)-GPS show that beaver dams are typically flooded on higher high tides, only impounding water at low tide to allow beaver movement during this time. Tidal beaver dam density per km was more than twice that reported in the fluvial literature, while mean dam head was about 80% and mean dam height about 60% of fluvial dams. Low-tide beaver pool depths were 75% of fluvial beaver ponds, while pool areas were 67% of their fluvial counterparts. The comparable beaver dam metrics between fluvial and tidal systems suggests their role in tidal ecosystems may be comparably significant. Inspection of Google Earth aerial photographs back to 1990 indicated that tidal beaver dams can persist for at least 35 years, spanning several generations of beaver. Given the ecosystem importance of beaver in fluvial and lacustrine habitats, better understanding of the distribution and ecosystem role of beaver in tidal wetland habitat, and the geometry of their dams and low-tide pools, would likely allow more effective restoration of estuarine habitat that is critical to a variety of fish and wildlife, including threatened species such as Chinook and coho salmon.

 SeaDoc Society A23-3415-S002 https://orcid.org/0000-0001-6286-7935 HoodW. Gregory This project was financially supported by the SeaDoc Society, a program of the Karen C. Drayer Wildlife Health Center, School of Veterinary Medicine, University of California, Davis. The award number was A23-3415-S002. Additional support was provided by the Tulalip Tribes, the Swinomish Indian Tribal Community, and the Sauk-Suiattle Tribe. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Data Availability All data files are available from Zenodo. The data DOI is: 10.5281/zenodo.18894036. Introduction

Beaver (Castor canadensis [North America] and C. fiber [Eurasia]) are iconic and charismatic animals, so known for their industrious construction of dams in streams that they have inspired the phrase, “busy as a beaver”. A result of their dam construction and excavation of small channels in floodplains is thorough alteration of their environment to benefit wetland and riparian vegetation which provide food for the beaver. Fish, birds, and other wildlife also benefit from the habitat changes that result from increased ponding along otherwise lotic systems [1–4]. Beaver dams and their ponds can profoundly alter fluvial hydrology, geomorphology, and ecosystem function [5]. For this reason, beaver are the archetype ecosystem engineer [6,7].

While the importance of beaver to fluvial ecosystems has long been known, the presence of beaver in tidal systems has been largely overlooked. In the Skagit Delta (Puget Sound, Washington, USA), where twice-daily tides can vary by four meters, intertidal beaver dams and lodges are common, but little is known about how beaver in tidal systems compare to those in more “traditional” fluvial systems. Yet there are indications that their influences on tidal ecosystems could be as significant as those on fluvial ecosystem. For example, recent work has shown that Skagit Delta beaver dams quadruple tidal channel (low-tide) pool habitat for fish, compared to tidal channels without beaver [8]. The Skagit beaver pools also contain triple the density (by volume) of threatened juvenile Chinook salmon (Oncorhynchus tshawytscha) compared to low-tide channel shallows. At low tide, channels without pools often dewater almost completely, forcing most fish to emigrate to larger, deeper channels with subtidal habitat. Channels with low-tide pools may allow (1) longer residence of small fish in productive tidal marsh channel habitat, (2) lower energy expenditure because they fish are not forced to move to other areas, (3) perhaps lower predation pressure because they are not forced into deeper waters on the ebb tide, and (4) perhaps greater food resources because low-tide beaver ponds appear to trap a lot of organic detritus [8], though these hypothesized benefits to fish have yet to be proven.

Tidal estuarine ecosystems have generally been considered an unexpected place to find beaver. The scientific literature on fluvial beaver is extensive, while for tidal beaver it is nearly non-existent. There are significant differences between tidal and fluvial systems that beaver must confront. The most prominent are bidirectional flow (flood vs. ebb tides) vs. unidirectional flow; typically twice daily water level fluctuations in systems that are micro-tidal (< 2 m fluctuations), meso-tidal (2–4 m), macro-tidal (4–6), or hyper-tidal (6–15 m), where low tides in most systems can drain many channels entirely, vs. seasonal and episodic floods or droughts (which can also occur in tidal systems fed by rivers); and often the presence of a range of salinities ranging from freshwater (< 0.5 ppt) to seawater (35 ppt) to hypersaline in arid areas, which contrasts with solely freshwater in fluvial systems. However, salinity stratification (freshwater floating on heavier saltwater) is common, so some estuarine plants and animals can reduce their exposure to salt stress. Tidal marshes tend to have significantly higher drainage density than uplands; their networks are frequently interconnected by loops so that they are reticulated rather than dendritic like fluvial networks; and tidal marshes are very flat so that drainage divides are often hard to define, while fluvial divides are characterized by significant topographic relief [9]. Altogether, these network differences make tidal systems very two-dimensional while fluvial systems are comparatively one-dimensional.

Because so little is known about beaver in tidal wetlands, knowledge gaps are extensive. Thus, the goal of this paper is to provide fundamental natural history information on intertidal beaver dams and to highlight the need for further investigation of beaver in tidal habitats. This paper is structured by two foci of investigation. The first focus is an exploratory analysis of local-scale patterns of intertidal beaver dam and lodge distributions in tidal wetlands of the Skagit and Snohomish deltas, the two largest rivers draining into Puget Sound (Washington, USA). The goal of this focus is to explore spatial patterns in dam and lodge locations (a) to provide insight into the abundance of beaver structures and their distribution relative to estuarine habitats, e.g., characterized by vegetation communities and channel networks; (b) to allow comparison with fluvial beaver, e.g., are there significant differences in dam or lodge density that might suggest differences in habitat quality or beaver behavior; (c) to understand the function of dams for beaver in tidal environments and how that compares with fluvial dams; and (d) to contribute guidance for tidal wetland restoration design that accommodates beaver and the ecosystem functions they may provide. The second focus is an assessment of the region-scale distribution of intertidal beaver dams in the Pacific Northwest, from British Columbia to Oregon, which shows that tidal beaver are widespread in the region. Region-scale comparisons are also needed to understand variation in beaver behavior that might be associated with system variation, e.g., large rivers with their greater freshwater discharge provide more freshwater tidal habitat, while small coastal wetlands with limited freshwater inflow may be more saline and thus more challenging habitat for beaver. Both foci support the argument that tidal wetlands are not marginal habitat for beaver, rather, they are a place where highly adaptable beaver can thrive. They are also places where we need to pay more attention to the ecosystem role of beaver.

 Methods Study areas

This study was focused on the tidal wetlands at the mouths of the two largest rivers in Puget Sound: the Skagit and Snohomish rivers, which respectively deliver 34% and 19% of the freshwater input to Puget Sound as a consequence of mean annual flows of 475 and 270 m3s-1 [10]. The Skagit River has a classic, triangular delta lying on top of extensive lahar deposits from volcanic eruptions 6000 and 2000 years ago that extended the river mouth 25 km into Puget Sound and created extensive, low, flat areas [11–13]. Most of the delta has been isolated from riverine and tidal influence by levees and dikes, and converted to agriculture and other uses. Consequently, the historically dynamic delta has been tamed by anthropogenic flood control so that geomorphic activity has been reduced and constrained to the two active sub-deltas at the termini of the two principal river distributaries (branches), the North Fork and South Fork, of the Skagit River where historical tidal marsh remains and where marsh progradation has since occurred [14]. The Snohomish estuary is comparatively narrow, 3–4 km wide and 20 km long, and confined by hills on both sides all the way to Puget Sound. Within this stretch, the river consists of anastomosing distributaries in a tidal floodplain dissected by blind-end tidal channels (henceforth, “tidal channels”).

Coastal systems of the Pacific Northwest are generally characterized by mixed semidiurnal tides, i.e., two high and two low tides per day of unequal range. The mean diurnal tide range (difference between higher high and lower low tides) in the Skagit and Snohomish deltas is 3.6 m, but this can vary geographically in Puget Sound from 2.6 m in the north to 4.6 m in the south [15]. Mature marsh surfaces are generally at the elevation of mean higher high water (MHHW), i.e., the average of the daily higher high tides, but during higher high spring tides, marsh surfaces can be inundated by up to 1 m of water, depending on marsh location and elevation. Salinity in coastal wetlands is inversely related to the amount of freshwater inflow and with distance from the mouth of the estuary. These salinity patterns influence vegetation zonation, with salt tolerant species typical of small coastal systems with limited freshwater inflows, and freshwater marsh and swamps (dominated by woody species) typical of systems with significant freshwater inflows or areas further inland along the estuarine salinity gradient.

Field surveys were focused on tidal shrub and Sitka spruce (Picea sitchensis) dominated zones of the Skagit and Snohomish deltas, because previous work [8] and local field experience showed that beaver dams and lodges were usually absent in tidal marshes dominated by herbaceous plants such as the sedges and bulrushes (Cyperaceae) that are typical of Pacific Northwest oligohaline tidal marshes. Tidal shrub habitat provides the woody building material that beaver need for dam and lodge construction. Thus, beaver infrastructure is rarely far from tidal shrub habitat. Tidal shrub and forest are found in three principal areas of the Snohomish Delta: the Quilceda marsh near the mouth of the river; Heron Point, about 5300 m upstream from the river mouth; and Otter Island, 5600 m upstream of the river mouth. These areas are separated by herbaceous marsh that were historically flood-breached and abandoned farmlands, and various intentional tidal marsh habitat restoration sites. The Quilceda marsh consists mostly of herbaceous marsh dominated by sedge (Carex lyngbyei), soft-stem bulrush (Schoenoplectus tabernaemontani), and non-native cattail (Typha angustifolia and T. x glauca) in increasing order of marsh elevation. At still higher elevations, and in a smaller portion of the marsh, typical shrubs such as willow (Salix spp.), black twinberry (Lonicera involucrata), and wild rose (Rosa nutkana) are found, followed finally by Sitka spruce. Heron Point consists mostly of Sitka spruce tidal swamp with an understory of shrubs, while Otter Island is mostly shrub dominated, especially wild rose, with about a quarter of the island comprised of Sitka spruce tidal swamp.

Compared to the Snohomish, the Skagit delta marshes in the North Fork and South Fork sub-deltas are a patchier mosaic of herbaceous and shrub vegetation, with scattered Sitka spruce trees. Shrubs and trees predominate along channel banks, but some channels can have one bank with woody vegetation and the other with herbaceous cover. Unlike the Snohomish, sweetgale (Myrica gale) is common in the Skagit marshes and forms dense, nearly impenetrable thickets. This aromatic shrub is shaded out by willows at slightly higher elevations. Wild rose is not as common in the Skagit as the Snohomish delta. More information on the shrub and tree communities in the Skagit can be found in [16].

Within the South Fork sub-delta there are several areas that have been restored from agricultural use to tidal wetlands to provide rearing habitat for juvenile Chinook salmon, a threatened species and an important regional food source and cultural icon. One of these areas, Milltown Island, had its protective dikes breached by a flood in the 1970s, and was subsequently abandoned and allowed to passively return to tidal wetland habitat. Because of its relatively high elevation in the tidal frame, willows have colonized significant portions of the site, as have beaver. Thus, beaver dams and lodges were surveyed on this site to compare with reference marshes in the Skagit and Snohomish deltas.

 RTK-GPS field surveys

Field surveys were conducted at low tide with a real-time kinematic (RTK)-GPS (3-cm horizontal and vertical resolution) in the Snohomish delta (October 2023; March, April, June 2024), the North Fork sub-delta (September 2023; April 2025), and Milltown Island (September 2020). Elevations were referenced to the North American Vertical Datum of 1988 (NAVD88). Tidal channels in the Snohomish and Skagit delta study areas were chosen for accessibility due to logistic and safety challenges working in a meso- to macro-tidal environment. Milltown Island, on the other hand, was almost completely censused except for a few areas where access was impossible due to deep water (even at low tide) and dense shrub thickets. Other dams and lodges were mapped in the South Fork sub-delta in an older effort using hip-chains and field photos [8]; the South Fork data set was periodically updated as incidental field observations during other work produced new sightings of intertidal beaver dams and lodges. RTK-GPS surveyed channel profiles included measurements on the channel bottom, the water surface at low tide, channel banks, the lowest top of beaver dams (where water would spill over), and the ground elevation and tops of beaver lodges (to estimate lodge height). Beaver scent mounds were rarely observed, but when they were, they were also located with RTK-GPS. ArcMap GIS was used to plot RTK-GPS data, generate channel profile traces, and produce maps of the field data.

 Statistical analysis

Simple linear regression was done using an on-line calculator [17]. Residuals were evaluated for deviations from normality by the Shapiro-Wilk test and examination of the residuals plot. Analysis of Covariance (ANCOVA) was done using an on-line calculator, with normality and homoscedasticity checked by the Shapiro-Wilk and Levine’s test, respectively [18].

G-tests of goodness of fit were done using the Statology calculator [19] and applied to comparison of the count of movement routes from observed beaver lodge locations vs. the locations of random points on channel banks. Random points were generated in GIS (ArcGIS Pro, ESRI) using the “Create Random Points” geoprocessing tool, applied independently to a shapefiles of the Skagit River distributaries and of Skagit Delta saltmarsh tidal channels. For each shapefile, 100 points were generated and the number of movement routes within 15 m and within 50 m were counted for each observed lodge and each random point. The observed lodge count for each category of route count was sometimes too low for G-test analysis, so this count was standardized by conversion to a percentage of the total number of observed lodges and multiplied by 100 for comparison with the 100 random points. Categories of route counts that still had too few observations for analysis were lumped together in logical groups to allow analysis.

 Results Local-scale field surveys

Intertidal beaver dams and lodges (S1-S9 Figs in S1 File) were surveyed in three locations in the Snohomish Delta. In the Quilceda marsh, near the mouth of the river, 41 intertidal dams and 3 lodges were encountered in a little over 2000 m of tidal channels in the portion of the marsh that was characterized by tidal shrub vegetation and Sitka spruce trees (Fig 1). Due to challenging field conditions only 67% of the terrain could be sampled, so total dam and lodge counts could be 50% higher than observed. At Heron Point the surveyed tidal channels were extensively dammed, and flooded understory channel-ponds ramified throughout. Thus, unsurveyed channel was similarly difficult to assess as for Quilceda, but at least 1000 m of channel could be detected in aerial photographs that were not surveyed for beaver. Thus, only 45% of the available tidal channel may have been surveyed, so that almost twice as many dams and lodges may be present here as observed. Only a portion of Otter Island is densely covered by Sitka spruce canopy; the rest is covered in a variety of water-tolerant shrubs. Thus, aerial photo inspection could identify approximately 2000 m of likely channel that was not surveyed, so total dam and lodge count could be as much as 2.4 times what was observed.

10.1371/journal.pone.0349313.g001Overview of beaver dam and lodge locations in the Snohomish estuary.

Yellow lines are traces of field-surveyed tidal channels; red dots are beaver dams; white triangles are beaver lodges; gray triangle is an abandoned lodge; white star is a scent mound. Areas without tidal shrub vegetation were not surveyed, because previous work had shown that beaver structures were absent there. Map Data © 2019 Google.

The North Fork and South Fork Skagit tidal marshes are a mosaic of shrub and herbaceous marsh, so total unsurveyed channel could be confidently estimated from aerial photographs. In the North Fork sub-delta, approximately 5100 m of likely channel were not surveyed, nearly half the amount that was surveyed (Fig 2), so total dam and lodge count could be 50% higher than observed. In the larger South Fork sub-delta, 28.7 km of tidal shrub channel were not surveyed compared to the 13.5 km that were surveyed (Fig 3), so total beaver dam and lodge count could be three times as high as observed. In the Milltown Island tidal habitat restoration site on the east side of the South Fork delta, channel surveys were exhaustive so that 12.0 km of channels were canvassed, while only 1.1 km could not be accessed due to dense shrub thickets and deep water. Thus, it is likely that only 9% of the site’s dams were missed.

10.1371/journal.pone.0349313.g002Overview of beaver dam and lodge locations in the North Fork Skagit sub-delta tidal wetlands.

Yellow lines are traces of field-surveyed tidal channels; red dots are beaver dams; white triangles are beaver lodges. Areas without tidal shrub vegetation were not surveyed, because previous work had shown that beaver structures were absent there. Heavily forested areas are non-tidal, rocky outcrops. River flow is from the east side of the frame. Farmland borders the study area to the north. Imagery © 2021 Eagleview.

 10.1371/journal.pone.0349313.g003Tidal beaver structures in the eastern half of the South Fork Skagit delta marshes.

Yellow lines are surveyed channel traces; red dots are beaver dams; white triangles are lodges. The Milltown Island restoration site is highlighted (blue oval). Imagery © 2021 Eagleview.

Dam density (per km of surveyed tidal channel) ranged from 10.3 to 29.6 km-1, and averaged 19.1 km-1 for the Snohomish and Skagit reference marshes (Table 1). In contrast the Milltown Island restoration site value was 6.0 km-1, less than 1/3 of the reference marsh average. Lodge density ranged from 1.4 to 3.7 km-1, and averaged 2.1 km-1. The value for Milltown Island was 0.25 km-1, which is less than 1/8th the reference marsh average. Beaver colonies can occupy more than one lodge, but this study did not distinguish between active versus inactive lodges. For fluvial and tidal colony densities to be comparable, tidal beaver would have to construct 3.4 lodges per colony (cf. S1 Table in S1 File).

10.1371/journal.pone.0349313.t001Summary statistics on estuarine tidal dam and lodge density.
Location Channel length surveyed (m) Count of dams Count of lodges Dams per km of channel Lodges per km of channel
Snohomish DeltaQuilceda marsh 2,010 41 3 20.4 1.5
Heron Point 810 24 3 29.6 3.7
Otter Island 1,400 18 2 12.9 1.4
North Fork Skagit delta 10,520 235 20 22.3 1.9
South Fork Skagit delta 13,450 139 29 10.3 2.2
Milltown Islanda 12,020 72 3 6.0 0.25
Tidal mean (sans Milltown Island) 19.1 2.1

aMilltown Island is abandoned farmland, whose dikes were breached by a flood in the 1970s.

In all surveys of the Skagit and Snohomish deltas, only one bank lodge was found; it was located on a large river distributary levee that was augmented in size by turn of the century (19th-20th) deposition of dredge spoils to allow ship navigation. All other beaver lodges were constructed on marsh surfaces and consisted of the classic mounds of sticks and mud. Mound lodges were invariably located on the banks of either tidal channels or river distributary channels. Of the 59 lodges found in the Skagit and Snohomish deltas, 16 were found on distributary banks and 43 on tidal channel banks. In the vicinity of the lodges, distributaries averaged 74 m wide (median = 42 m, range = 7–350 m) while tidal channels averaged 3.1 m wide (median = 2.6 m, range = 0.7–19 m).

Lodges on distributary banks had at least two aquatic movement routes from the lodges, upstream and downstream, but they were often located near distributary junctions or tributary tidal channels, which increased beaver movement options in the lodge vicinity. Lodges located within tidal channel networks were typically located in the vicinity (within 15 m or within 50 m) of tributary junctions, often multiple junctions (Fig 4), which may be a way of maximizing beaver movement routes away from the lodge. Comparison of observed lodge locations with random points suggests that beaver bias the location of their lodges towards sites with high connectivity in channel networks, i.e., the number of potential aquatic movement routes. Their lodges had significantly higher connectivity than points randomly located along channel network banks (Fig 4). This was true at two spatial scales: within 15 m of a lodge (or random point) and within 50 m, and it was true for lodges located on distributary banks as well as tidal channel banks. Tidal channel lodges had much higher connectivity than distributary bank lodges, especially at the 50-m scale, which is consistent with most lodges being located on tidal channels. However, the presence of a large fraction, 27%, of all lodges on distributary banks suggests there may possibly be some compensatory benefit to location on a distributary.

10.1371/journal.pone.0349313.g004Frequency distributions of the number of aquatic movement routes in the vicinity (< 15 m and < 50 m) of beaver lodges vs. random points located on tidal channel or distributary banks. Channel profiles

Beaver-dammed tidal channel profiles show striking contrasts between downstream channel reaches that flow through herbaceous marsh and upstream reaches that flow through tidal shrub and Sitka spruce vegetation. The downstream reaches have no beaver dams for 300–400 m, so at low tide there are no pools, and the water is typically 9–15 cm deep while flowing over relatively narrow (< 1-m wide) thalwegs at low tide; the broader (4–8 m wide) channel bottoms outside of the thalweg are essentially drained at low tide. The upstream reaches over the next 300–400 m have a series of five to seven dams that pond water in a stair-step fashion so that the water surface at low tide eventually attains the channel bank top. The most seaward dams are clearly overtopped by flood tides, but the highest, landward dams are likely only overtopped on higher-high spring tides (compare dam height vs. mean higher high water [MHHW] in Fig 5). At low tide, beaver pond depths in the tidal channels typically range from 30 to 95 cm (x¯ = 50.6, n = 23). Similar profiles can be found in the South Fork Skagit Delta marshes when downstream tidal channel reaches are bereft of tidal shrub vegetation and instead are characterized by typical Pacific Northwest sedges.

10.1371/journal.pone.0349313.g005Typical beaver-dammed tidal channel profiles.

Water surface elevations are at low tide; compare with mean higher high water (MHHW) [top graphs]. Associated channel traces (yellow lines) are shown over a 2019 air photograph with beaver dams (red dots) plotted [bottom photo]. Other channel trace profile points are not shown to avoid graphic clutter. White triangles are beaver lodges; the small gray triangle is the remains of a former lodge; the white star is a scent mound. Areas further upstream were not mapped because they were diffusely anastomosing, pond-like channels that were difficult to map. Map Data © 2019 Google.

Beaver lodges (S9 Fig in S1 File) were not always located where dams raised low-tide water levels to near the marsh surface, but they were generally located where dams raised the low-tide water level sufficiently to cover the lodge entrances. However, the purpose of tidal beaver dams is not merely to flood lodge entrances, it is also to allow beaver to swim in tidal channels during low tide. This is illustrated by dam construction in two senescing river distributaries in the North Fork Skagit delta where lodges are not present (Fig 6). The North Fork mainstem channel has avulsed (starting in 2004) into a new course that bypasses most of the delta [20]. The avulsion channel has become the new mainstem channel of the North Fork, while the former mainstem channel and its distributaries are filling with sediment and shoaling. Prior to shoaling the distributaries hosted no beaver dams, likely because they were deep enough to contain substantial subtidal waters that allowed low-tide beaver movement, and because their large size allowed tidal flows or river floods strong enough to wash away beaver dams. As the distributaries have shoaled, they have also narrowed and filled with vegetation, but the aggrading vegetated surface is still lower than the adjacent, older marsh platform. Thus, these shoaling distributaries do not host any beaver lodges, neither on their current low-elevation banks (because the lodges would be flooded at high tide), nor on the nearby older, pre-avulsion distributary banks (one such pre-existing lodge was abandoned because it became land-locked). Because the shoaling distributaries drain in two directions at low tide, the distributary channel profiles are convex upward, and the beaver dams create low-tide pools that never stair-step the water surface elevation up to MHHW, the approximate elevation of a mature marsh platform. The only possible function of these dams for beaver is to create pools that allow low-tide beaver movement.

10.1371/journal.pone.0349313.g006Profiles of two senescing delta distributaries.

Surveyed channel traces (yellow lines) are superimposed on a 2021 aerial photo (Imagery © 2021 Eagleview); red dots are beaver dams [top photo]. Insert is 2004 photo prior to distributary senescence (Imagery © 2004 Triathlon). Surveyed channel profiles [bottom graphs] show low-tide conditions. The mature marsh surface is at approximately MHHW (e.g., the first profile point on the top graph), while the marsh surface in the shoaling distributaries is lower. For more examples of beaver channel profiles see S10-S13 Figs in S1 File.

Intertidal beaver dams were always constructed of mostly sticks, from nearby shrubs, and supplemented with mud. Sometimes bank slumps were incorporated into dams, suggesting beaver opportunistically taking advantage of such situations to reduce construction effort. On one occasion a derelict boat that filled a tidal channel was incorporated into a large beaver dam. Dam heights ranged from 5 to 172 cm and averaged 48 cm (Table 2). This average is a little over half the average height reported for fluvial dams (Table S2 in S1 File). Water head for tidal beaver dams ranged from 0 cm for dams in disrepair to 134 cm, with an average of 13 cm. If dams in disrepair are ignored, then the average water head is 25 cm, which is 78% of the average reported for fluvial dams. Intertidal beaver dam pool depths ranged from 4 to 137 cm and averaged 51 cm, which is 75% of the average for fluvial beaver dam pools. Intertidal beaver dam pools are generally linear in form, i.e., tidal channels that are not allowed to drain on low tide. But some dam pools flood beyond the banks of the tidal channels into adjacent topographic lows, so they are more irregular in shape. There is likely some negative bias in the tidal beaver dam pools size estimates because they were measured in GIS from air photos, so canopy cover prevented measurement of some pools, and very narrow channels were a challenge for distinguishing pools. Reported fluvial beaver dam pool areas averaged about 50% larger than tidal beaver dam pools.

10.1371/journal.pone.0349313.t002Beaver dam and pool metrics for tidal systems. Mean (standard deviation) and range of tidal beaver dam water heads and heights, and pool depths and areas, as measured from surveyed tidal channel profiles and GIS analysis of aerial photographs. <italic>N</italic> is the sample size for each metric, respectively. Fluvial weighted means from Table S1 in <xref ref-type="supplementary-material" rid="pone.0349313.s001">S1 File</xref> are shown for comparison.
Site N Water head (cm) Dam height (cm) Pool depth (cm) Pool area (m2)
Snohomish Delta– Quilceda 24, 27, 27, 21 36 (19)10–85 57 (24)21–114 47 (21)15 - 95 70 (73)10 - 356
Snohomish Delta– Heron Point 19, 21, 18, 8 45 (37)8 - 127 71 (31)23–141 68 (23)26 - 105 49 (33)21 - 109
Snohomish Delta– Otter Island 17, 19, 13, 15 34 (34)7 - 134 73 (36)23 - 189 70 (22)21 - 101 251 (173)24 - 540
Skagit Delta– North Fork 143, 223, 216, 62 19 (12)7 - 63 43 (26)5 - 143 49 (25)4 - 137 120 (165)3 - 874
Tidal weighted meana 25 48 51 123
Fluvial weighted meanb 32 82.3 68 184

aWeighted by sample size; bSee Table S1 in S1 File for more details.

It is reasonable to think that channel size could affect beaver dam construction. If we have a simplistic hypothesis that assumes tidal dams are generally relatively uniform in design (with some variation around a mean), then each dam would have a similar height and water head, so that a nearly constant number of dams would be found in tidal channels regardless of channel size (length), because the difference between low tide channel bottom elevations and MHHW is similar for almost all channels (except small tributaries to tidal channels). If true, then short channels should have steeper gradient and closely spaced dams while long tidal channels should have lower gradient and more widely spaced dams.

Exploratory statistical analysis indicated that the spacing of beaver dams was not uniform or random, but instead increased with channel length (Fig 7), consistent with the hypothesis. The linear regression relationship was calculated for the fourteen North Fork Skagit Delta tidal channels (R2 = 0.85, p < 0.000004) and the six Snohomish tidal channels (R2 = 0.71, p < 0.035), with ANCOVA showing no significant difference in dam spacing between the two groups (F1, 17 = 0.96, p = 0.34), but significant influence of the covariate, channel length (F1, 17 = 55.2, p < 0.001). Points for the senescing North Fork distributary channels plotted in the vicinity of the two regression lines.

10.1371/journal.pone.0349313.g007Exploratory regression analysis of beaver dam spacing and dam head relative to channel length and gradient for North Fork (NF) Skagit tidal channels, NF senescing distributary channels, and Snohomish tidal channels.

In addition to channel length, channel water surface gradient might also correlate with beaver dam spacing, so gradient was compared to channel length. The two Heron Point channels (Snohomish Delta) were distinct outliers from all other channels, with gradients four times the average of the other channels. For the other channels, channel length was inversely related to gradient, as expected, but only explained 20% of the variance in gradient (R2 = 0.20, F1,19 = 4.89, p = 0.04). Nevertheless, dam spacing was not related to channel gradient (R2 = 0.14, F1,19 = 2.93, p = 0.10), although the trend appeared consistent with the hypothesis.

Because the Quilceda Marsh (Snohomish) channels had similar gradients as the North Fork Skagit channels, they were no longer distinguished in subsequent analyses, but Heron Point channels continued to be unexpected outliers. Channel length and mean dam spacing each affected mean dam head (R2 = 0.60, F1,16 = 23.9, p = 0.0002 and R2 = 0.44, F1,16 = 12.5, p = 0.003, respectively). The dams in the senescing distributaries were also outliers, but this was unsurprising given the immature morphology of these channels, whose banks were well below MHHW. Maximum dam head showed similar relationships to channel length and mean dam spacing, indicating that dams are not generally uniform as assumed in the hypothesis, but instead respond to channel size. Muddier sediments in the Snohomish Delta compared to the sandier Skagit, and on the convex bank of a Snohomish distributary meander (Heron Point) may allow construction of taller and less leaky dams in the Heron Point marshes.

 Region-scale aerial photo surveys

Aerial photo inspection to identify likely locations of beaver dams and lodges could allow planning of focused field work to maximize beneficial effort. To this end, aerial photo

interpretations of likely dam and lodge locations were compared to field observations, and the degree of agreement was summarized in error matrices (Table 3). Only dam presences were predicted from aerial photo interpretation, not dam absences, to avoid inflating estimates of prediction accuracy. Thus, the error matrices are missing the quadrant containing verified negative predictions.

10.1371/journal.pone.0349313.t003Error matrices for aerial photo interpretation of tidal beaver dams. “Predictions” are aerial photo interpretations; “Observations” are ground-truthed.
North Fork Skagit Delta
Dam observed Dam not observed Total
Dam predicted 69 52 121
Dam not predicted 197 –
Total 266
Snohomish Delta
Dam observed Dam not observed Total
Dam predicted 31 20 51
Dam not predicted 57 –
Total 88
Fraser, Nisqually, Chehalis, Grays Harbor, Willapa Bay, Oregon Coast
Dam observed Dam not observed Total
Dam predicted 43 1 44
Dam not predicted 14 –
Total 57

True dams were 57% of all predicted dams for the North Fork Skagit Delta and 60% for the Snohomish Delta. Predicted dams were 26% of all field-observed dams for the North Fork Skagit Delta and 35% for the Snohomish Delta, which means that when using aerial photos, I identified only 1/4–1/3 of the dams that were actually present on the landscape. The sources of error in photo interpretation stem primarily from the presence of shrub canopies that obscure channels, location of small dams in narrow, steep-sided channels that makes their observation difficult, and general photo ambiguity (Table 4). Photo ambiguity generally consisted of difficulty in distinguishing a dam from an accumulation of wrack or sticks in a tidal channel, or of distinguishing a dam from a local high spot in the channel bottom that has been exposed by low tide.

10.1371/journal.pone.0349313.t004Sources of photo interpretation error for tidal beaver dam prediction. “Success” = true positive; “Fail” = false positive; “Missed dam” = false negative.
North Fork Skagit Delta Success Fail Missed dam TOTAL
Ambiguity 31 39 84 154
Shrub canopy 5 45 50
Small channel 4 64 68
Classic impoundment 29 29
Visible dam 5 5
TOTAL 65 48 193 306
Snohomish Delta Success Fail Missed dam TOTAL
Ambiguity 19 18 17 54
Shrub canopy 1 34 35
Small channel 1 1 6 8
Classic impoundment 11 11
Visible dam
TOTAL 31 20 57 108

Successfully predicted dams were most confidently identified by clearly visible dams and by photo-signatures I termed “classic impoundments”, which consisted of a combination of a channel obstruction that demarked a narrow downstream channel from a strikingly wider upstream channel (Fig 8). When the dam was the first in a series, the downstream channel often had clearly exposed substrate at low tide, while the upstream channel was clearly inundated.

10.1371/journal.pone.0349313.g008Examples of photo interpretation of intertidal beaver dam locations.

All photos are at low tide. Yellow arrows indicate correctly predicted dams; black arrows indicate missed dams, i.e., not predicted, but observed in the field. Photos A and B are examples from the Quilceda marsh in the Snohomish estuary, where ebb-tide flow is from top to bottom in each photo (Map Data © 2022 Google). Photo C is from the North Fork Skagit sub-delta, where ebb-tide flow is from right to left (Imagery © 2023 Eagleview).

The lessons learned from the error analysis were applied to a search of Google Earth imagery for intertidal dams in other coastal wetlands in the Pacific Northwest to determine how widespread the distribution of tidal beaver might be. Google Earth photos from 2012 to 2024 (earlier photos generally had insufficient resolution to be useful) were examined for reliable indicators of tidal beaver dams. Ambiguous photo-signatures, canopy covered areas, and narrow channels were ignored in favor of clearly observable dams and classic impoundments. Intertidal beaver dam photo-signatures were observed from the Fraser River delta near Vancouver, Canada to the Siuslaw River estuary, Oregon (Fig 9), where mean diurnal tidal range varies from 2.6 to 4.6 m in the Salish Sea, and from 2.1 to 3.1 m on the outer coasts of Washington and Oregon. Ground-truthing of a sample of these conservatively identified dams revealed much lower rates of false positives, i.e., 43 out of 44 photo-based predictions were verified in the field (the one error was due to anthropogenic widening of a short channel reach), but 14 of 57 dams observed in the field were still missed in photo examination (Table 3).

10.1371/journal.pone.0349313.g009General locations of confirmed and likely presence of intertidal beaver dams in the Pacific Northwest (red crosses).

See Supplementary Materials for more specific locations.

These coastal systems can be classified into two broad geomorphological categories: (1) large river systems with substantial freshwater input (e.g., Fraser, Nooksack, Skagit, and Snohomish deltas; Chehalis, Willapa, Columbia, Gray’s, Young’s, Salmon, and Siletz river estuaries), and (2) small coastal streams with modest freshwater input. The structure of beaver dam networks in large river systems has already been described earlier using the Skagit and Snohomish deltas as examples, but in small coastal streams the beaver dam network is sometimes different. There, a few ordinary channel-spanning tidal dams may precede a large, ½-m to 1-m high, marsh-spanning dam that impounds a large pond on the marsh surface (Fig 10). These marsh surface dams ranged from 40 m to 120 m long and impounded ponds that ranged from 400 m2 to 6000 m2 in surface area. The ponds typically had a small tributary creek that provided freshwater inflow.

10.1371/journal.pone.0349313.g010Examples of marsh surface dams (red lines) and impoundments for small tidal stream systems.

Channel spanning dams are noted by yellow push-pins in low-tide Google Earth imagery. Redman Slough and Red Salmon Creek were ground-truthed. For scale, the Redman slough surface dam was 65 m long; the longer Red Salmon Creek surface dam was 120 m long. Map Data © 2016 Google.

Being able to confidently identify tidal beaver dams in aerial photos allows estimation of minimum dam age from available historical aerial photos, where the estimate depends on image quality. Examination of aerial photos shows that Pacific Northwest tidal beaver dams can easily reach a couple of decades in age; in a couple of locations, they are at least 35 years old (Table 5).

10.1371/journal.pone.0349313.t005Estimated minimum tidal beaver dam age from earliest observation in an aerial photograph.
Location Earliest observation Latest absence Minimum age
Fraser Delta 2004 – 21
Snohomish-Quilceda 2003 1990 22
Snohomish-Otter Isl. 2010 – 15
Skagit North Forka 2004 – 21
Nisqually 2005 2003 20
Chehalis 2006 – 19
East Fork Redman Slough 1990 – 35
Johns River trail sloughs 1990 – 35
Elk River Estuary 2011 – 14
Columbia Estuary 2012 – 13
Youngs River 2004 – 21

“Latest absence” refers to the oldest photograph in which the beaver dam is not observed. Dam age is thus bounded by earliest observation and latest absence. Latest absence is not noted when photo quality was too poor to reliably indicate that dam was present or absent.

aAn associated lodge was also clearly visible and present since 2004.

Lodges are nearly impossible to identify from aerial photographs because they are typically either obscured by shrub canopy, or their coloration differs little from background vegetation. They were successfully predicted only three times, out of 45 lodges found in the Skagit tidal wetlands and 8 found in the Snohomish tidal wetlands, on the three rare occasions when there was a suspicious contrast in coloration.

 Discussion

Beaver have been considered a riverine and lacustrine species, but recent observations indicate they can also inhabit estuarine systems with significant tidal ranges (up to 5 m). This paper clearly documents the presence of beaver in tidal wetlands from the Fraser Delta in southern British Columbia to the Siuslaw estuary midway along the Oregon coast. This is likely not just a west coast phenomenon as beaver have also been observed in tidal wetlands of the Tivoli Bays in the Hudson River National Estuarine Research Reserve (New York) where there is a 1.7-m tidal range (Jack Loud, Bard College student, personal communication; S38 Fig in S1 File).

Tidal beaver dams and lodges are abundant in large river deltas and estuaries such as the Snohomish, Skagit, Fraser (S14-S16 Figs in S1 File), Chehalis (S19-S22 Figs in S1 File), and Columbia (S33 Fig in S1 File), which have abundant freshwater discharge, low marsh salinities (fresh to oligohaline), and abundant woody vegetation in the tidal zone. In these systems, tidal beaver dams and lodges are found in close association with tidal shrub and tidal forest vegetation. More surprisingly, beaver can also be found in smaller coastal systems with sparse patches of woody vegetation or woody vegetation along the upland margins of the tidal marsh. To find woody material to build a dam, beaver have to travel as far as 70 m, 74 m, and 200 m from a tidal dam to a forested border for Netarts Bay (S34-S35 Figs in S1 File), the Siletz estuary (S36-S37 Figs in S1 File), and a marsh along the Johns River (S23-S24 Figs in S1 File), respectively, although almost all of that distance can be traversed by tidal channels. In contrast to larger river deltas or estuaries with high freshwater inflows and large coastal freshwater plumes, beaver in small coastal tidal systems may have limited scope in downstream movement because of high downstream salinities. To compensate, beaver in small, coastal, tidally-influenced streams with small freshwater inflows may impound large ponds with marsh surface dams to provide greater amounts of freshwater habitat. The in-channel dams downstream of the marsh-surface dam may not only assist in damming the upstream pond, they may also serve as salinity barriers by blocking the densest, saltiest tidal floodwaters. While small coastal systems may have a relatively small amount of freshwater input, inland portions of the systems may be stratified, with lighter freshwater floating on top of denser saltwater, so that beaver experience minimal salt stress. Vegetation observed in the more landward portions of these small coastal systems was invariably typical of fresh to oligohaline tidal marshes.

Tidal habitat use by beaver may appear surprising given more common observations in fluvial systems, but this perception may be a consequence of range contraction following near extinction of the species by 19th century trapping, as well as greater habitat loss in estuarine ecosystems (e.g., 85% loss on the west coast [21]) compared to riverine and lacustrine systems. Tidal shrub and forest wetlands, preferred tidal beaver habitat, have been disproportionately lost compared to tidal emergent marsh [22]. Compared to tidal marsh, these areas are also more challenging to human movement and are thus less studied. Given the abundance of beaver dams and lodges in tidal wetlands throughout the Pacific Northwest, and the abundance of food resources for beaver in these wetlands, it seems unlikely that tidal wetlands are sub-optimal beaver habitat compared to “normal” fluvial habitat. They are simply additional, but different habitat. However, conclusively demonstrating this will require collecting data on body condition and breeding statistics for tidal beaver.

The role of tidal beaver dams

From the perspective of a beaver, the function of a dam is to provide water deep enough to inundate lodge entrances and allow beaver to safely swim to foraging sites [23]. This is true in fluvial systems and appears to also be the case in tidal systems, except that in tidal systems the need for inundation is focused on low tides. During high tides, tidal channels and most beaver dams are completely inundated. Indeed, the marsh surface itself can be flooded by up to a meter, depending on the tide, weather, and location in the estuarine elevation gradient. Thus, during high tides, beaver dams are generally irrelevant. However, during low tides, beaver dams shine. Channels without beaver dams often convey shallow streams of water, as little as only a few centimeters deep. In contrast, channels with beaver dams have low-tide pools averaging 51 cm deep, enough for beaver to swim at low tide. Because low-tide beaver dams can quadruple the length of low-tide pools compared to channels without beaver dams (other pools can be formed by scour in meander cut banks, at tributary junctions, and around logs [8]), their role in increasing beaver mobility at low tide appears to be significant. This is starkly illustrated in the profiles of the Quilceda tidal channels, where the downstream half of the channels have no beaver dams and convey a relative trickle of water, while the upstream half is impounded by a series of dams that provide almost continuous swimming routes, interrupted only by the dams themselves.

Contrasts between the Quilceda channel profiles and those of the senescing North Fork Skagit distributaries raise the question of beaver mobility, especially during low tide. All of the Quilceda channel profiles are without dams in their lower half, and consequently have water depths of only a few centimeters there. This suggests that beaver residing in this area have relatively restricted movements, at least during low tides, moving around only in the upper half of the channel networks where dams pond water at low tide. The North Fork Skagit channel profiles, on the other hand, have dams down to or near the channel outlets, and the senescing distributaries have dams well distant from any lodges or tidal channel networks. This suggests that beaver in the North Fork delta are potentially more mobile during low tides, traveling farther from their lodges and traveling through river distributaries, even those that are senescing and shoaling. Why the apparent difference between Quilceda and the North Fork Skagit? Habitat does not appear to be limiting; there is an abundance of vegetation to eat in the North Fork delta, especially willows. Are population densities higher in the North Fork? Or are the Quilceda marshes distinctive because they are located so close to the river mouth and experience relatively high salinity? During summer low river flows the salinity in this area averages 18.7 ppt, while Heron Point and Otter Island average 9.4 and 3.8 ppt, respectively [24]. The Skagit marsh areas that host beaver are typically 0–5 ppt during summer low flows. Perhaps the high late-summer salinities discourage Quilceda beaver from venturing far from their Sitka spruce strongholds and produce the more restricted distributions of their dams.

Beaver dams in fluvial systems have ecosystem roles beyond those intended by beaver; plant and wildlife species richness increase as a result of beaver dam ponding, and fluvial beaver dams can more generally affect stream hydrology, geomorphology, and ecosystem function [5]. Tidal beaver dams increase low-tide, volumetric fish density by 3- to 7-fold depending on species, indicating that tidal beaver ponds may provide important low-tide refuge for many fish [8]. In the current study, dam functions were not evaluated, but incidental observations suggest at least some dams, typically those higher in the channel network, can accumulate significant amounts of unconsolidated sediment. Walking through those dam ponds was hazardous, with soft sediments a meter or more in depth. Several small, headward channel reaches in the North Fork delta were so full of semi-consolidated sediment that mud reached to within a half meter of the marsh surface and the channels appeared to be in the process of senescing to the point of disappearance as a result of sediment trapping by the beaver dams. Tidal beaver dams appear to trap abundant organic detritus as well as mineral sediments, and this may result in high production of invertebrate detritivores [8]. Low-tide beaver ponds may also increase the residence time of juvenile salmon and other fish in the channel networks; and predation on fish in these ponds may be relatively low. Finally, a provocative thought is that beaver dams might serve as salinity barriers that reduce saltwater intrusion into tidal channel networks. Because saltwater is heavier than freshwater, a series of beaver dams might sequentially freshen tidal flood waters along a tidal channel profile. Perhaps this is why beaver, and Sitka spruce, can inhabit the brackish Quilceda marsh. Clearly, there is scope for a great deal of further investigation into the possible ecosystem roles of tidal beaver dams and their low-tide ponds.

 Single large or several small beaver dams

In many tidal channels, a series of beaver dams stair-steps water up to marsh surface to create extensive areas of constant ponding, even during low tides. Why does stair-stepping occur? Why not just build one tall dam that ponds up water throughout the tidal channel? This option was observed, but only once, on the east side of Otter Island in the Snohomish Delta where a nearly 2-m tall beaver dam was located 20 m from the channel outlet. At low tide, the water head for the dam was 1.38 m, forming an 87-cm deep pool upstream of the dam, compared to a trickle of water downstream (the difference between dam head and pool depth suggesting perhaps 51 cm of sediment trapping in the pool). More commonly, intertidal beaver dams occur in a series and average 48 cm tall with water heads averaging 25 cm (i.e., the water depth on the downstream face of the dam averages 23 cm). Water head is related to pressure or force on the dam, and is proportional to seepage through the dam [25–27]. For dams generally, whether man-made, formed by landslides, or beaver built, seepage refers to leakage of water through the dam or its foundation. Excessive seepage can cause internal erosion within the dam body or erosion of its foundations, which weakens the dam’s structure and can lead to failure. Seepage can be controlled by injecting grout (mud for beaver) into permeable areas to fill voids and reduce permeability, and by blanketing the dam face on its upstream side with impermeable material (mud for beaver dams). Consequently, high water head beaver dams likely have higher risk of failure than low-head dams and likely require more maintenance by beaver to reduce this risk. These risks are likely exacerbated by building on soft sediments typical of muddy to sandy tidal channels. Stair-stepping reduces the hydraulic head for an individual dam thereby reducing seepage, instability, and maintenance. A single large dam also leads to greater risk in catastrophic loss of beaver habitat in a channel compared to a series of dams which provides some redundancy that results in only partial loss of beaver habitat with a single dam failure. The nearly 2-m high dam that was observed, was only 24 m distant from a beaver lodge, the proximity perhaps facilitating frequent dam maintenance.

 Habitat restoration

Puget Sound Chinook salmon and Oregon Coast coho salmon (O. kisutch) are threatened species under the Endangered Species Act. Because juveniles of both species benefit significantly from rearing in tidal marsh habitats [28], there has been considerable investment in the region for tidal marsh habitat restoration to recover these species. With the recognition that beaver are a natural part of tidal wetlands in the Pacific Northwest, and likely elsewhere, and given their status as ecosystem engineers, their role in ecosystem restoration and recovery should be investigated [8]. Tidal marsh restoration can potentially benefit beaver as well as juvenile salmon and other fish and wildlife, as long as the habitat needs of beaver are considered, i.e., sufficient supply of freshwater, tidal channels, shrubs, trees, and herbaceous food plants. Beaver colonization of the Milltown Island restoration site in the South Fork Skagit Delta shows that restoration sites can support beaver, though the lower density of beaver dams and lodges on the site, compared to reference marsh conditions, suggests thoughtful consideration of beaver habitat needs could improve beaver responses to restoration projects. Beaver occupancy of habitat restoration sites could in turn benefit juvenile salmon rearing on the site [8].

Further research on how beaver adapt to tidal marsh conditions and how their dams and low-tide pools interact with other biota and eco-geomorphic processes is necessary before rashly rushing to translocate beaver to tidal marsh systems. It is important, for example, to understand what the beaver carrying capacity of different types of tidal marshes might be, and whether beaver are already pushing those limits in any given location. Where do dispersing estuarine juveniles go once they are ready to establish their own territories? Are they more likely to migrate along the coast as they search for territorial opportunities, or move inland? How do we account for beaver landscape connectivity in the course of tidal marsh restoration so that juvenile beaver have opportunities for dispersal?

Tidal marsh restoration planners and engineers in the Pacific Northwest (e.g., the Washington Department of Fish and Wildlife and the Columbia River Estuary Task Force) have shown interest in constructing beaver dam analogs (BDAs [29,30]) in tidal channels. However, there are no design guidelines for BDAs in tidal systems (though this paper provides data that could be used for this purpose), and it is unclear whether they are appropriate for tidal systems. In fluvial systems, BDAs are typically installed in incised streams to trap sediment and raise the stream bed to restore floodplain connectivity. The geomorphic role of tidal beaver dams is still unknown, and their ecological role still needs better resolution, so it seems premature to incautiously adapt BDAs to tidal channels. Uncritically applying fluvial paradigms to tidal systems is likely unwise given the many differences between fluvial and tidal systems, particularly bidirectional flow, salinity stratification, and twice-daily fluctuations of 1–5 m in water surface elevation in tidal systems. There is a great deal of tidal beaver natural history that needs to be understood before we can wisely proceed with beaver reintroductions or BDA installations in tidal marsh restoration sites.

 Summary

In the Pacific Northwest, at least, beaver are not unusual in tidal wetlands that contain or are bordered by woody vegetation. Tidal beaver are widespread and occur in a wide variety of coastal settings, reflecting the adaptability of beaver and the likelihood that they are important ecosystem engineers in estuarine as well as riverine systems.

In tidal systems, beaver build at least two types of dams. The most common type of dam is located in tidal channels and is composed of sticks and mud, sometimes incorporating bank slumps and sometimes overgrown with herbaceous wetland vegetation, such as sedge. Typically, these dams range in size from 1 to 4 m wide and from 20 to 100 cm tall, though larger dams are sometimes encountered. The dams typically stair-step low tide water levels up to the marsh surface providing extensive areas of low-tide refuge for beaver and other tidal channel denizens. During high tides most, if not all, of the beaver dams are completely inundated so that movement by tidal channel fauna, such as fish, is unimpeded. The dams are therefor functional mostly at low tide, serving to allow beaver movement for foraging and to permanently inundate lodge entrances.

The other kind of dam is much less common. It is located on marsh surfaces, oriented perpendicularly to a tidal channel, and includes a dam within the channel. This surficial dam ranges from 0.5 to 1.0 m in height, but can be several hundred meters long to produce large ponds on the marsh surface. These dams are found principally in small coastal systems with limited freshwater inflow from small streams. The function of the dams appears to be to trap the freshwater and augment the amount of freshwater habitat available to the resident beaver.

The widespread occurrence of beaver in low-salinity tidal wetlands that allow easy access to herbaceous and woody vegetation for forage and building materials suggests that this habitat is perfectly suitable for beaver, rather than sub-optimal compared to better-documented fluvial beaver systems. It also suggests that neglecting the role of beaver in coastal wetlands will lead to an incomplete understanding of these systems and deficient management, including for habitat restoration and protection.

 Supporting information Tables S1-S2; Figs S1-S38.

(DOCX)

Thank you to Joe Gaydos, SeaDoc Society, for his enthusiastic support for this project. Thanks to Todd Zackey, Tulalip Tribes Department of Natural Resources, for assistance in surveying the Snohomish Delta tidal channels. Thanks to Skagit River System Cooperatve biologists who assisted in RTK-GPS surveys in the North Fork Skagit Delta: Kathleen McKeegan, Neal Robertson, and Dennis Newman. Thank you to Eric Balke, Ducks Unlimited Canada, for photos and GPS waypoints of beaver dams that he observed in the Fraser Delta tidal marshes. Finally, thank you to Jack Loud who contacted me about the beaver lodges he saw in the Hudson River National Estuarine Research Reserve and who sent me lodge photographs and documentation of GPS waypoints.

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7 Jan 2026

-->PONE-D-25-54787-->-->Beaver in tidal habitat: Examples from the Pacific Northwest-->-->PLOS One

Dear Dr. Hood,

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Reviewer #1: Partly

Reviewer #2: Yes

Reviewer #3: Partly

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Reviewer #1: Yes

Reviewer #2: Yes

Reviewer #3: Yes

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Reviewer #1: No

Reviewer #2: Yes

Reviewer #3: No

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Reviewer #2: Yes

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-->5. Review Comments to the Author

Please use the space provided to explain your answers to the questions above. You may also include additional comments for the author, including concerns about dual publication, research ethics, or publication ethics. (Please upload your review as an attachment if it exceeds 20,000 characters)-->

Reviewer #1: This is a very interesting journal article where the author demonstrates significant knowledge about the system and includes an impressive amount of fieldwork, documenting beavers pushing into marine habitats, something that can significantly extend the knowledge and understanding of beavers. I want to congratulate the author on recognizing this pattern and aiming to publish this important contribution!

Unfortunately, in its current form, the manuscript suffers from a lack of clear research questions, clarity in the result and method section and fails to highlight the potential implications of this paper (which are definitely there). I think with some work, especially on introduction, presentation of the results and discussion, these limitations can be addressed, and this paper can fulfil its full potential to an international readership.

Especially crucial is to rework the result section. Full of jargon and site names, I got lost a lot trying to understand where we are and what is being talked about. For a primarily freshwater ecologist (which most beaver people are), this was challenging, and marine terms and context need a much better introduction.

In my opinion, the author has to decide where this story should be published and in what format. If it is to be published in an internation journal with the aim to put the idea out that beavers also live in tidal habitats, then the manuscript needs to be trimmed and generalized for the internation reader. If the author would like to tell a more detailed story including discussing site specifics, then a local naturalist journal might be the better choice.

Data availability

Where is the data available? Please be more specific, ideally with a link.

Abstract

Line 22 Please spell out or remove the abbreviation RTK

Line 24 Please mention density per km

Introduction

General comments

Nice introduction with very good sections. Some parts can be shortened/condensed (BDA’s, part on fish from Line 51). Given this has the ambition to be a naturalist paper informing the international readership about the extraordinary capabilities of beavers extending into tidal marches, I would like some more description of the habitat and beaver biology compared to the more standard beaver freshwater stream habitat. Some of that information may be in the methods already, I would consider moving it. This will help the reader understand why beaver colonize these habitats and ultimately also how that could be applied to other areas.

Additionally, from the current introduction, the research questions are not so clear to me and that could be achieved with a better structuring of the introduction clearly highlighting knowledge gaps.

Finally, I miss an overview or at least the question of other places where beavers push into saltwater/marine habitats. Is this know from other places than the Pacific Northwest? I think some literature is available. I think also beavers in Argentina were known to colonize islands separated by saltwater and feed on marine food like algae. Check Anderson et al. 2008 Mammal review doi: 10.1111/j.1365-2907.2008.00136.x as a starter.

Line 35 please specify where each beaver is native (broadly) and which beaver species is dealt with in this manuscript

Line 60 I am not sure why suddenly the text is about BDA’s so much. I understand that there seems to be a management/knowledge gap, but I am not sure the specific examples (maybe limit to one) add much for the international reader. Maybe that is also something for the discussion?

Line 65 As mentioned in the comment below, I would expand a little bit the physical differences of tidal ecosystems to more linear stream ecosystems. Maybe also worth to expand more generally about differences in biology of the beaver (why is it in the tidal marshes, what is it building dams out of, what is it feeding on)

Line 70 From here on, I got confused. The first sentence is clear, but then I feel the last sentence might be more appropriate and I also do not understand why the small stream are inhospitable.

Maybe a broader introduction into the dynamics of tidal habitats could help follow the reader here. The sentence from line 71 I would move to the end and start with something like “to do this, we choose the following study regions representing ….”

Methods

Whatever you decide to do with the results, follow the same logic in the methods.

I think a simple map, or at least a better description where these sites are and how much they are separated would help the reader. I am not even sure the country has been mentioned. I am generally confused by the many localities mentioned and specific histories that I am not sure are important for the paper’s questions.

Line 139 Spell out abbreviation for RTK

Results

The result need a lot of restructuring. Currently, this is not publishable, as it is hard to follow and often lost in details. I tried to suggest improvements, because I think the manuscript idea is great!

I suggest 3-4 Figures, in total one per section.

Whitin the sections:

- Separate your results better. Answer the main questions.

- A lot of the site-specifics has to go to help the reader better follow. Consider to elaborate and describe site specifics in the appendix. The results need to be generalized (at least the text).

Maybe have two section types. First a section of the dams and transects (things you measured), then a section about naturalist history, eg how the dams are build (things you observed).

I think some of these Figure S1-S9 can go into a composite figure in the main manuscript to illustrate the habitat.

In turn, I would reduce the figures in the main manuscript (or do a composite figure) for Figures 1-6

Line 160 I do not understand how this 2000 m were selected, at random, is this everything or accessibility? This should be mentioned in the methods. Again I am a bit at loss which name refers to which area and a map and a clear structure/separation in methods results per area could definitely help. Also for the other areas, data about the size belongs to methods in my opinion.

Line 162 to line 170 I would just mention in the results that due to challenging field conditions only half of the terrain could be sampled. The speculation on line 170 belongs into the discussion, same with line 174 and line 179.

Line 213 I am sorry, but I cannot follow this sentence. What is lodges per colony? Not even “colony” has been introduced as a term. While this might be clear for beaver people, it could create confusion for other readers. Also what does it has to do with Table 2?

Line 220 I suggest to move this table to supplementary materials. I like that you back up your claims with data, but would generally advice to reduce numbers of figures and tables in the main story to focus on the most important contributions.

Line 223 First it was not clear to me that now lodges are reported. Maybe here (and in other places) separate sub-headings could help to guide the reader?

Line 235 It is not entirely clear to me what a blind or a distributary channel is, this has not been introduced. Best place probably in the methods. Same with movement routes, this not be properly defined. What movement is it? Water, beaver? What is a route count?

Line 246 Maybe it helps to add to larger sampling region (here Snohomish) so the reader keep some simpler overview of the sites

Line 255 What is the mean higher high water? Keep in mind, beaver people are often not very knowledgeable in marine jargon, you need to explain these things much better. Probably I miss some terms as well in this review, check again that you explained all terms very well. At this point I think a glossary could be helpful!

Additionally: What is navd88? Is it needed in the figure legend?

Also clearly label which graph belongs to which line on the figure

Apply these changes to all figures, of course.

Line 269 Why is there two red dots in the figure shown when it mentions: Other channel trace profile points are not shown to avoid graphic clutter

Line 280, from “probably”: This is discussion up until the “provides an example” Also again from 297. I currently do not understand what the added value of the extensive description of this (seemingly) separate case study is. What does the reader learn that is important for the main story? Maybe some of this natural history and curiosity (which has its value!) can go in the description of the Fig S9?

Line 327 I would separate the table and pit the literature results for freshwater into the appendix, and summarize them in text. Why has N different numbers, is this the sample size per “stream”?

Section starting from Line 340: Always use p < 0.001 if p-value is very small

Line 358 This could be the best figure of your manuscript, but currently I do not understand what is going on. Why is Snohomisch differet to NF blind (what is NF) and NF distrib? What are the different regression lines for and why do the y differ from the ones in the text? I would suggest: Simplify the figure, show one regression per figure (overall). Show site-specific regressions in the appendix and condition specific regression only on the main text if they are significant.

Line 370 I find it hard to understand to follow this error matrix. Can you add a sentence that helps how to read it? For example: “reading example” In the North Fork area we predicted 121 dams, whereas we observed 69…” I would also move this table to the appendix and add an “overall” category

Line 387 That is partly not results, but discussion

Line 393: I would suggest t move that into the appendix

Line 399 That is partly methods. I think you do not mention this part in the methods at all?

Line 422 That is a very cool result and overview!

Line 425 That is discussion from here

Line 442 Another candidate for the supplementary materials

Line 458 That is the sort of detail that can be probably safely removed from the manuscript without loosing too much knowledge. Same with the paragraph 460 onwards (maybe one sentence in the discussion)

Discussion

In general, the discussion is too long, often too far away from what the actual research topic was (a lot of speculation and open questions not backed up by the data) and lacking integration into other research. A lot of interesting points are there, but buried in the text. Again I would emphasize not to use site names (only exceptionally), but instead generalize. If people are interested in specific sites, they can still contact the author, but for most international readers the local site names do not matter.

Line 493 Interesting, can you elaborate a bit more what you mean with freshwater habitat?

Line 526 Lot’s of individual site names again in this paragraph, maybe it is possible to reduce these? Here you are trying to generalize, not to discuss individual cases. Maybe saying “some sites have high molbility, some low” is therefore sufficient.

Line 535 I would avoid questions and frame it as research gaps: “It is unclear why”

Line 573 Again, I would avoid questions and frame it as research gaps: “It is unclear why”

Line 598 For me that whole paragraph is speculation and has nothing to do with the presented data (no sound measurments done, not introduced in the introduction). It is interesting, but for me needs another study. Maybe keep 1-2 sentences.

Line 625 For me, also these parts are discussed to much being to far from the research topic. If I am not mistake the increased fish densities were mentioned earlier already, so maybe there 1-2 points can be taken up

Reviewer #2: This is a well written piece and very interesting topic, though potentially to a limited audience, it still supports further knowledge expansion regarding a highly popular species of conservation and active restoration interest. I suggest minor revisions to improve readability.

Personally, suggest any rewriting of any first-person statements eg lines 53-55

Any further exploration of Eurasian beavers in tidal zones, we certainly witness this but agree little documentation

Line 57 reintroducing rather than introducing for correct conservation biology term - surely beavers here historically

Detailed methodology and a range of variables discussed though I think I’m missing more on the wider beaver population density and history on when how why beavers have entered this system, I guess we presume it is population density pressure pushing beavers to live in theoretically more challenging and ‘sub optimal’ areas - are you seeing this as a result of this - therefore would releases into such areas (as briefly proposed) without other habitat infilling just result in released animals moving inland to ‘easier’ habitat and therefore release site fidelity would be challenging for conservation projects and a nice link for BDA investigations as pointed out - perhaps expand this point? Personally, I’m interested in how well are the beavers doing in these areas - any insight into body conditions, breeding stats etc?

Good use of tables to summarise results as some of the text can be a bit dense to read and take in - I just wonder is there any scope to put any more of the local scale field survey results into a table - just a question - like habitat/ forage type access?

Suggest there is repetition eg use of aerial photos in the regional scale section - could be more concise here along with discussion lines 545-569 - please consider condensing

Discussion - sound of water - I would consider this inclusion entirely - there is plenty of evidence that this is more of a repeated fable rather than accepted scientific and field experience, I am pained to give more light to this and would question what it brings to this otherwise well considered study. A quick survey of many beaver dams proves they have the sound of running water - and if not this can change over the course of a day / week or weather condition and many other damming triggering behaviours are widely reported in the scientific literature

Line 634 paragraph - feel this is the place to discuss the practicalities and animal welfare considerations of releasing beavers into such regions, wider population density, release site fidelity etc

Reviewer #3: This ms presents interesting data on the form and distribution of beaver dams in tidal habitats. Some of the data is descriptive. However, overall I found the paper to lack focus and therefore it was difficult to follow at times. Probably what is needed is for a tight set of aims (perhaps with hypotheses, but not necessarily so), with the results and discussion structured around these. I also couldn’t find the data on OSF despite the author saying it was there, so this needs to be checked. I think the data could benefit from some additional statistical modelling to better understand the pattern of dam building and lodge construction in these environments. There is a good dataset here that deserves to be used. Some of the discussion near the end could be tightened up too. I’ve put a few specific comments below. I hope this gets resubmitted as there is very little in the literature on the subject of beavers occupying tidal areas.

Lines 47-51. My immediate question here is whether tidal beaver dams act as a fish trap during receding tides, rather than being ‘chosen’ by fish as refuges during low tide. Clearly, these juvenile salmon like these tidal channels, otherwise they wouldn’t move up there in the first place. But, though out with the scope of this study, understanding the effects of these dams on juvenile salmon habitat selection, mortality, growth rates and other life history characteristics would be very interesting. My main point in commenting on this is that unless one conducts the above research, we can’t be certain these beaver pools ‘support’ triple the density rather than just happen to hold this greater density as a result of ‘entrapment’ of individuals moving in and out of the creeks with the tidal fluctuations, so I suggest using the word ‘hold’ in place of ‘support’ in line 48.

Line 139. Please provide the full name on first mention of RTK-GPS.

Lines 138-148. Please provide the dates (years and months) of the various parts of the field work.

Lines 220-222 (Table 2). I don’t recall any description of how the literature review was conducted. Is this representative sample of literature on dam frequency? I also find it interesting that the damming frequency is lower in fluvial and lacustrine environments (and no individual study as high as this one), because I imagine the range of stream gradients would be higher away from intertidal areas. But then again, perhaps intertidal dams are smaller (maybe due to constraints from the bidirectional flow) and so more are needed. I hope you discuss this later.

Lines 226-233. Ideally one would compare mound lodge locations with a random set of bank-side points and us a binomial model to analyse whether the selection of the various channel types (blind, distributary or main channel?) and distance to distributary junctions / confluence was statistically significant.

Lines 231-233 and Fig 4. Was the difference in frequency distribution statistically significant. I think a non-parametric test comparing these (e.g. KS or Mann-Whitney U) would be possible here.

Table 3. As my comment on Table 2.

Lines 335-340. Curious. Were shorter channels also of higher gradient? How did the water heads compare in shorter and longer channels?

Table 4. I find this initially a little confusing, so I think this needs a short explanation in the table caption or change in the column/row labels. ‘Predicted’ means identified or otherwise from aerial photographs and ‘observed’ is ‘ground-truthed’ data.

Table 5. This is also confusing, despite the footnote (which I initially missed anyway). Could you change the wording? Success is a true positive? Fail is a false positive? Missed dam is a false negative?

Line 530. I think clearer wording needed: not all readers might understand colloquialisms such as “homebodies”. Perhaps just say they have reduced home ranges.

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Reviewer #1: No

Reviewer #2: No

Reviewer #3: Yes:  Ruairidh D. Campbell

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 10.1371/journal.pone.0349313.r002 Author response to Decision Letter 1 Submission Version 1

6 Mar 2026

My responses to the editor and reviewers are collected in the attached file "Response to Reviewers". I would only add that I appreciate the constructive criticism, and the positive feedback and encouragement from the reviewers.

It would be easiest to look at the above referenced docx file, but it you want all of my responses included here, I have copied and pasted the docx text below. The copied text is not rich text so it misses out on bolding and text color that I use to distinguish reviewers’ and editors comments from my responses.

4. We note that Figures 1, 2, 3, 5, 6, 7, 9, 10, 11, S1, S10, S11, S12, S13, S14, S15, S16, S17, S18, S19, S20, S21, S22, S23, S24, S25, S26, S2, S28, S29, S30, S31, S32, S33, S34, S36, S37, and S39 in your submission contain [map/satellite] images which may be copyrighted. All PLOS content is published under the Creative Commons Attribution License (CC BY 4.0), which means that the manuscript, images, and Supporting Information files will be freely available online, and any third party is permitted to access, download, copy, distribute, and use these materials in any way, even commercially, with proper attribution. For these reasons, we cannot publish previously copyrighted maps or satellite images created using proprietary data, such as Google software (Google Maps, Street View, and Earth). For more information, see our copyright guidelines: http://journals.plos.org/plosone/s/licenses-and-copyright.

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Author’s Response:

I have consulted the copyright acknowledgement requirements for Google, ESRI, and Pictometry (Eagleview Inc.), the sources the aerial imagery in my figures. None of them require requests for permission and Google specifically says that they do not have time to respond to such requests. Instead, they provide clear guidance for when their imagery can be use and how to acknowledge their copyright. I have detailed my response regarding content permissions in an attached “Other” file for each source of imagery.

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Reviewers’ comments:

Reviewer’s Responses to Questions

Comments to the Author

1. Is the manuscript technically sound, and do the data support the conclusions?

The manuscript must describe a technically sound piece of scientific research with data that supports the conclusions. Experiments must have been conducted rigorously, with appropriate controls, replication, and sample sizes. The conclusions must be drawn appropriately based on the data presented.

Reviewer #1: Partly

Reviewer #2: Yes

Reviewer #3: Partly

________________________________________

2. Has the statistical analysis been performed appropriately and rigorously?

Reviewer #1: Yes

Reviewer #2: Yes

Reviewer #3: Yes

________________________________________

3. Have the authors made all data underlying the findings in their manuscript fully available?

The PLOS Data policy requires authors to make all data underlying the findings described in their manuscript fully available without restriction, with rare exception (please refer to the Data Availability Statement in the manuscript PDF file). The data should be provided as part of the manuscript or its supporting information, or deposited to a public repository. For example, in addition to summary statistics, the data points behind means, medians and variance measures should be available. If there are restrictions on publicly sharing data—e.g. participant privacy or use of data from a third party—those must be specified.

Reviewer #1: No

Reviewer #2: Yes

Reviewer #3: No

________________________________________

4. Is the manuscript presented in an intelligible fashion and written in standard English?

Reviewer #1: No

Reviewer #2: Yes

Reviewer #3: Yes

________________________________________

5. Review Comments to the Author

Please use the space provided to explain your answers to the questions above. You may also include additional comments for the author, including concerns about dual publication, research ethics, or publication ethics. (Please upload your review as an attachment if it exceeds 20,000 characters)

Reviewer #1:

This is a very interesting journal article where the author demonstrates significant knowledge about the system and includes an impressive amount of fieldwork, documenting beavers pushing into marine habitats, something that can significantly extend the knowledge and understanding of beavers. I want to congratulate the author on recognizing this pattern and aiming to publish this important contribution!

Unfortunately, in its current form, the manuscript suffers from a lack of clear research questions, clarity in the result and method section and fails to highlight the potential implications of this paper (which are definitely there). I think with some work, especially on introduction, presentation of the results and discussion, these limitations can be addressed, and this paper can fulfil its full potential to an international readership.

RESPONSE: I have clarified to research objectives of the paper in the Introduction and I have added a Summary after the Discussion to clarify the “take-home” messages of the paper and highlight the implications of the paper.

Especially crucial is to rework the result section. Full of jargon and site names, I got lost a lot trying to understand where we are and what is being talked about. For a primarily freshwater ecologist (which most beaver people are), this was challenging, and marine terms and context need a much better introduction.

RESPONSE: I added a paragraph in the Introduction and in the Study Areas section of the Methods to provide a brief primer on tidal systems, including necessary jargon. Unfortunately, this jargon is essential vocabulary for estuarine ecology.

In my opinion, the author has to decide where this story should be published and in what format. If it is to be published in an internation journal with the aim to put the idea out that beavers also live in tidal habitats, then the manuscript needs to be trimmed and generalized for the internation reader. If the author would like to tell a more detailed story including discussing site specifics, then a local naturalist journal might be the better choice.

Abstract

Line 22 Please spell out or remove the abbreviation RTK

RESPONSE: I spelled out the abbreviation.

Line 24 Please mention density per km

RESPONSE: Changed as suggested.

Introduction

General comments

Nice introduction with very good sections. Some parts can be shortened/condensed (BDA’s, part on fish from Line 51). Given this has the ambition to be a naturalist paper informing the international readership about the extraordinary capabilities of beavers extending into tidal marches, I would like some more description of the habitat and beaver biology compared to the more standard beaver freshwater stream habitat. Some of that information may be in the methods already, I would consider moving it. This will help the reader understand why beaver colonize these habitats and ultimately also how that could be applied to other areas.

RESPONSE: I added a paragraph to the Introduction that summarizes the principal differences between fluvial and tidal systems that beaver, and beaver biologists, must confront. I think this will be useful for beaver biologists who are typically unfamiliar with tidal ecosystems.

Additionally, from the current introduction, the research questions are not so clear to me and that could be achieved with a better structuring of the introduction clearly highlighting knowledge gaps.

RESPONSE: The issue of greater clarity in the goals of the paper was also mentioned by reviewer #3. I have edited the Introduction to clarify the structure of the paper and the goals of each structural component. I have also edited the Discussion to circle back on the goals described in the Introduction. With regard to highlighting knowledge gaps, there is almost nothing known about tidal beaver—almost everything is a knowledge gap (which I point out in the new edits), but I hope I have made it clear that I am addressing two particular gaps and pointing out the need for greater attention to tidal beaver in general.

Finally, I miss an overview or at least the question of other places where beavers push into saltwater/marine habitats. Is this know from other places than the Pacific Northwest? I think some literature is available. I think also beavers in Argentina were known to colonize islands separated by saltwater and feed on marine food like algae. Check Anderson et al. 2008 Mammal review doi: 10.1111/j.1365-2907.2008.00136.x as a starter.

RESPONSE: Yes, this story is well known. Beaver can swim through salty seas to disperse to islands or along shorelines. This has also been observed in Puget Sound. But dispersal is temporary transit. Occupation of estuarine (brackish) habitat is long-term and a much different phenomenon.

Line 35 please specify where each beaver is native (broadly) and which beaver species is dealt with in this manuscript.

RESPONSE: Parenthetical notes were made for each species at this point in the text. Given that the fieldwork in this manuscript takes place in North America, it did not seem necessary to explicitly say that this paper focused solely on C. canadensis—also there are literature-based comparisons of lodges and dams in a couple of tables that include both species, so to say that this paper is exclusively about C. canadensis is not precisely accurate. Both species have a lot in common, which is why I included C. fiber in the literature-based comparisons.

Line 60 I am not sure why suddenly the text is about BDA’s so much. I understand that there seems to be a management/knowledge gap, but I am not sure the specific examples (maybe limit to one) add much for the international reader. Maybe that is also something for the discussion?

RESPONSE: I have moved this text to the Discussion section dealing with habitat restoration. I have also edited it slightly to be more concise.

Line 65 As mentioned in the comment below, I would expand a little bit the physical differences of tidal ecosystems to more linear stream ecosystems. Maybe also worth to expand more generally about differences in biology of the beaver (why is it in the tidal marshes, what is it building dams out of, what is it feeding on)

RESPONSE: Line 65 (and this whole paragraph in general) has been moved to the Discussion, following the recommendation of another reviewer. I have also added a paragraph to the Introduction that describes the physical differences between fluvial and tidal systems, including differences in channel network geometry. However, with regard to differences in the biology of beaver between the two systems, nothing is actually known about these differences—there is no literature on this subject. The purpose of this paper is to explore some of these questions of tidal beaver biology. Thus, rather than discussing this in the Introduction, I think it more appropriate to do so in the Discussion.

Line 70 From here on, I got confused. The first sentence is clear, but then I feel the last sentence might be more appropriate and I also do not understand why the small stream are inhospitable.

RESPONSE: I am glad to see that the reviewer liked both sentences. I think they bracket the paragraph well. However, I revised the last part of the last sentence to clarify why small streams are inhospitable (low inflow = high salinity).

Maybe a broader introduction into the dynamics of tidal habitats could help follow the reader here. The sentence from line 71 I would move to the end and start with something like “to do this, we choose the following study regions representing ….”

RESPONSE: I added this recommended detail to the second paragraph of the “Study areas” section of the methods, where I discuss tides and salinity.

Methods

Whatever you decide to do with the results, follow the same logic in the methods.

I think a simple map, or at least a better description where these sites are and how much they are separated would help the reader.

RESPONSE: Maps of this kind was presented in Figs 1-3 (Fig 1 was revised to include the local-scale map in response to another comment) and Fig 9 in the results section (the reviewer liked Fig 9 a lot). It is challenging to keep figures to a minimum and avoid redundancy.

I am not even sure the country has been mentioned.

RESPONSE: A parenthetical statement was added to address this in the 1st line of the “Study Areas” section of the Methods.

I am generally confused by the many localities mentioned and specific histories that I am not sure are important for the paper’s questions.

RESPONSE: I have deleted mention of many of these localities in response to this reviewer’s comment and to reviewer #2’s recommendation that I be more “concise”.

Line 139 Spell out abbreviation for RTK

RESPONSE: Reviewer #3 made the same comment. Changed as suggested.

Results

The result need a lot of restructuring. Currently, this is not publishable, as it is hard to follow and often lost in details. I tried to suggest improvements, because I think the manuscript idea is great!

RESPONSE: In response to all three reviewers, I have edited the text to be more concise, remove unessential details (like the abundance of place names), and to focus on principal points.

I suggest 3-4 Figures, in total one per section.

RESPONSE: I have reduced the number of figures, but not to the extreme suggested by this reviewer.

Whitin the sections:

- Separate your results better. Answer the main questions.

- A lot of the site-specifics has to go to help the reader better follow. Consider to elaborate and describe site specifics in the appendix. The results need to be generalized (at least the text).

RESPONSE: In response to all three reviewers, I have edited the text to be more concise, and I have removed mention of place names as much as possible.

Maybe have two section types. First a section of the dams and transects (things you measured), then a section about naturalist history, eg how the dams are build (things you observed).

RESPONSE: I don’t think this distinction between measurement and observation is very clear or helpful. Measurements are a type of observation.

I think some of these Figure S1-S9 can go into a composite figure in the main manuscript to illustrate the habitat.

RESPONSE: I weighed this suggestion very carefully, because I love figures, but I also understand the need to be concise and focused. The Supporting Information document is a wonderful compromise although not every reader takes advantage of it. I don’t think even figures S1-9 do justice to the variety of tidal environments in which beaver dams can be found; all of the other figures in the Supplemental I

Submitted filename: Response to Reviewers.docx

 10.1371/journal.pone.0349313.r003 Decision Letter 1 Laruelle Goulven G Academic Editor 2026 Goulven G Laruelle This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Submission Version 1

14 Apr 2026

-->PONE-D-25-54787R1-->-->Beaver in tidal habitat: Examples from the Pacific Northwest-->-->PLOS One

Dear Dr. Hood,

Thank you for submitting your manuscript to PLOS ONE. After careful consideration, we feel that it has merit but does not fully meet PLOS ONE’s publication criteria as it currently stands. Therefore, we invite you to submit a revised version of the manuscript that addresses the points raised during the review process.-->-->

As you can see, both reviewers were largely satisfied with the corrections you did to the manuscript and convinced it now is close to a state in which it could be published. One of them nevertheless asks for a clarification regarding figure 7 and what the notations NF blind, NF distrib stand for and the other one, while acknowledging that the manuscript is very well written, suggestions that the editing could still be a bit improved, especially but making the research goals clearer.

Not being a native English speaker myself, I feel a little awkward asking one to improved his writing, especially because I also enjoyed reading the manuscript that I do find very well written indeed. However I also agree that some minor editing and attention to keep the flow of the text focused on the research question might help "holding the hand" of the reader along this really nice study.

I thus call this minor revisions because the option "superficial modifications" does not exist in the system. I will be very happy to accept the next version of your manuscript without sending it back for review if you agree to take into consideration these small requests.

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Reviewer #1: Thanks to the authors for a good revision, I think the manuscript has improved significantly, and I think it makes a valuable contribution to beaver and tidal science.

It was a lot easier to read now, and I appreciate the authors trying to better explain the different terms.

one small comment:

Figure 7: It is still not easy to understand what is NF blind, NF distrib. The abbreviation NF is not explained elsewhere in the manuscript as far as I saw. I assume it is North Fork. Maybe that can be put in the figure legend.

Reviewer #2: This is an epic piece of work, well written and informative and novel. I just find it could still benefit with robust editing as it is very long and detailed that I’m not sure improves the readability of the text eg vegetation descriptions, I’m getting a bit lost with the relevance to the study, which i still feel could have clearer research goals

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Reviewer #2: No

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 10.1371/journal.pone.0349313.r004 Author response to Decision Letter 2 Submission Version 2

22 Apr 2026

Responses to Reviewers and Editor (part 2) 22 April 2026

As you can see, both reviewers were largely satisfied with the corrections you did to the manuscript and convinced it now is close to a state in which it could be published. One of them nevertheless asks for a

[Reviewer] clarification regarding figure 7 and what the notations NF blind, NF distrib stand for

I made two changes to address this recommendation: [1] I added a parenthetical “(NF)” after the text “North Fork” in the manuscript figure legend (see track changes) to show that “North Fork” was being abbreviated to “NF”, and [2] I edited the legend box in the top figure changing “blind” to “tidal” to be consistent with the rest of the text, as the term “blind channel” was changed to merely “tidal” channel in response to previous recommendations by reviewers—I simply missed making the change in this figure; I also fully spelled out “distributary” to replace to previous abbreviation “distrib”.

and the other one, while acknowledging that the manuscript is very well written, [Reviewer] suggestion that the editing could still be a bit improved, especially by making the research goals clearer.

[Editor] I also agree that some minor editing and attention to keep the flow of the text focused on the research question might help "holding the hand" of the reader along this really nice study.

I have addressed these concerns by making edits in two locations in the manuscript. The first section of new text occurs from line 84 to 91 and consists of the following new text (underlined and blue) with original text in black and mostly removed. I believe this first section more explicitly details the goals of the paper, as requested.

“The goal of this focus is to explore spatial patterns in dam and lodge locations (a) to provide insight into the abundance of beaver structures and their distribution relative to estuarine habitats, e.g., characterized by vegetation communities and channel networks; (b) to allow comparison with fluvial beaver, e.g., are there significant differences in dam or lodge density that might suggest differences in habitat quality or beaver behavior; (c) to understand the function of dams for beaver in tidal environments and how that compares with fluvial dams; and (d) to contribute guidance for tidal wetland restoration design that accommodates beaver and the ecosystem functions they may provide. infer the role of beaver dams in tidal systems, from the perspective of beaver, and contrast it with their role in fluvial systems. An additional contrast is how beaver choose lodge locations in a two-dimensional estuarine landscape compared to a mostly one-dimensional riverine landscape.”

In a second location, from line 367 to 374, I added a paragraph (see text below) that describes a hypothesis that provides context for the results described in subsequent paragraphs and in Fig. 7. In three locations in the results, I also added phrases that indicated consistency or inconsistency with the hypothesis.

“It is reasonable to think that channel size could affect beaver dam construction. If we have a simplistic hypothesis that assumes tidal dams are generally relatively uniform in design (with some variation around a mean), then each dam would have a similar height and water head, so that a nearly constant number of dams would be found in tidal channels regardless of channel size (length), because the difference between low tide channel bottom elevations and MHHW is similar for almost all channels (except small tributaries to tidal channels). If true, then short channels should have steeper gradient and closely spaced dams while long tidal channels should have lower gradient and more widely spaced dams.”

Submitted filename: Response to Reviewers 22Apr2026.docx

 10.1371/journal.pone.0349313.r005 Decision Letter 2 Laruelle Goulven G Academic Editor 2026 Goulven G Laruelle This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Submission Version 2

29 Apr 2026

Beaver in tidal habitat: Examples from the Pacific Northwest

PONE-D-25-54787R2

Dear Dr. Hood,

We’re pleased to inform you that your manuscript has been judged scientifically suitable for publication and will be formally accepted for publication once it meets all outstanding technical requirements.

Within one week, you’ll receive an e-mail detailing the required amendments. When these have been addressed, you’ll receive a formal acceptance letter and your manuscript will be scheduled for publication.

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Kind regards,

Goulven G Laruelle

Academic Editor

PLOS One

Additional Editor Comments (optional):

As I said in my previous message, the final round of revisions was very minor and I appreciate that you took the time to address them carefully. I am now very happy to accept your manuscript as it now stands and forward it to the production team.

Reviewers’ comments:

 10.1371/journal.pone.0349313.r006 Acceptance letter Laruelle Goulven G Academic Editor 2026 Goulven G Laruelle This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

PONE-D-25-54787R2

PLOS One

Dear Dr. Hood,

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