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Article

Restored and Natural Wetland Small Mammal Communities in West Virginia, USA

by
Krista L. Noe
1,*,
Christopher T. Rota
1,
Mack W. Frantz
2 and
James T. Anderson
3
1
School of Natural Resources, West Virginia University, 1145 Evansdale Drive, Morgantown, WV 26506, USA
2
West Virginia Division of Natural Resources, Natural Heritage Program, 1110 Railroad Street, Farmington, WV 26571, USA
3
James C. Kennedy Waterfowl and Wetlands Conservation Center, Belle W. Baruch Institute of Coastal Ecology and Forest Science, Clemson University, P.O. Box 596, Georgetown, SC 29442, USA
*
Author to whom correspondence should be addressed.
Land 2022, 11(9), 1482; https://0-doi-org.brum.beds.ac.uk/10.3390/land11091482
Submission received: 30 July 2022 / Revised: 30 August 2022 / Accepted: 2 September 2022 / Published: 4 September 2022
(This article belongs to the Special Issue Wetland Construction and Restoration: Design and Performance)
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Abstract

:
Wetland restoration is a common practice, and, in many cases, it is for mitigation to offset losses of natural wetlands due to human interference. Researchers commonly compare bird, amphibian, and reptile communities between these wetlands and natural wetlands but overlook small mammals. However, terrestrial small mammals are essential to consider as they serve a fundamental role in the ecosystem as seed dispersers and prey for larger wildlife. We conducted small mammal trapping on 26 wetlands (n = 14 restored, n = 12 natural) in West Virginia, USA, in the summers of 2020 and 2021 to obtain and compare community metrics between wetland types. We found that mass, occupancy probability, and community composition were similar between restored and natural wetlands. However, the apparent abundance of deer mice (Peromyscus maniculatus) was higher in natural wetlands (p < 0.001). Because we captured the three rarest species exclusively in natural wetlands, the ability of restored wetlands to provide an adequate habitat for rare or wetland-obligate species may be biologically significant. Restored wetlands mainly offer sufficient habitat for small mammal communities, but apparent abundance in restored wetlands may differ from natural wetlands depending on species.

1. Introduction

Wetlands perform many functions, such as flood mitigation and improving water quality [1]. Due to the diverse functionality wetlands provide, many programs to restore wetlands exist. Many United States-based restoration programs are voluntary and run through the government, such as the Agricultural Conservation Easement Program [2] or the Partners for Fish and Wildlife Program [3]. There are also voluntary, non-governmental organizations that restore wetlands, such as Ducks Unlimited. Although many voluntary programs exist, wetland mitigation is compulsory; it is the lawfully imposed solution to planned human-induced wetland losses in the United States [4]. Wetland mitigation offsets losses to natural wetlands due to human activities, most commonly by creating or restoring wetlands [5]. Wetland creation involves the establishment of wetlands where they have not historically existed, and restoration involves reestablishing a wetland that humans previously drained or filled [6]. Researchers need to assess restored wetland functionality because of the widespread practice of wetland restoration and the assumption that these wetlands can truly replace natural wetlands [7,8].
One function of wetlands is to provide a habitat for wildlife. Wildlife responses to wetland conservation vary depending on taxa. Passerine occupancy probability, species richness [9], and total avian abundance [10] were similar between restored and reference wetlands in West Virginia, USA. Occupancy of some amphibian species was similar between natural and mitigated wetlands because restored wetlands provide chorusing [11] and breeding [12] habitats. However, these wetlands may not be viable for some amphibian species because some created wetlands dry up prematurely, leading to high larvae mortality and little recruitment [13]. The abundance of common turtle species was similar between restored and natural wetlands [14]. No differences in bat activity were observed between restored or created and natural wetlands, except in eastern pipistrelles (Perimyotis subflavus), which were positively associated with natural wetlands [15]. Furthermore, macroinvertebrate assemblages, a prey base for some wildlife species, were similar between natural and mitigated wetlands [16,17].
One largely understudied taxon in wetland restoration is terrestrial small mammals. The lack of data is problematic because terrestrial small mammals, such as rodents and shrews, play an essential role in the ecosystem. Terrestrial small mammals are significant seed dispersers, influencing the landscape [18]. Additionally, differing individual tendencies of scatter-hoarding species, such as deer mice (Peromyscus maniculatus), southern red-backed voles (Myodes gapperi), and northern short-tailed shrews (Blarina brevicauda), can affect how far seeds travel [19]. Aside from their importance as seed dispersers, they are important prey species for birds [20], bobcats (Lynx rufus) [21], and coyotes (Canis latrans) [22]. Terrestrial small mammals can also influence plant biomass, richness, and diversity [23]. Apart from their roles as seed dispersers, prey, and vegetation influencers, terrestrial small mammals can also be valuable bioindicators of restoration success or potential emerging human–wildlife conflict because of their sensitivity to changes in habitat [24,25,26]. The vegetative community impacts terrestrial small mammal diversity and abundance [27,28].
Researchers have not completely overlooked small mammals in restored wetlands. Species composition within the small mammal community varied with fluctuations in the hydroperiod and water depth in a restored wetland in Arkansas, USA [29]. The small mammal community was similar between planted and control areas in a South Carolina, USA, riparian wetland [30]. Kurz et al. [31] found small to mid-sized mammal abundance, richness, and species composition were similar between reference and restored wetlands in Ohio, USA. Existing research suggests small mammal communities respond to wetland restoration favorably. Age of restoration in these studies may have impacted the findings. Restored wetlands were no older than seven years at the time of research for Kurz et al. [31] and even less for Whitsitt and Tappe [29] and Wike et al. [30]. Researchers should sample a broad range of ages because restored wetlands become more like natural wetlands with age [32].
The objective of this study was to examine small mammal community characteristics in natural versus restored wetlands to determine if these wetlands contain similar small mammal communities. Wetland mitigation is designed to offset losses of destroyed natural wetlands. If restored wetlands, which account for many mitigated wetlands, cannot support similar community characteristics of vital wildlife taxa, then the wetland has not fulfilled its purpose in its entirety. Specifically, we examined (1) apparent abundance, (2) mass, (3) occupancy probability, and (4) community composition of small mammals between restored and natural wetlands. Given the finding by Balcombe et al. [32] that mitigation wetlands had higher vegetation species diversity, coupled with the reliance of small mammals on vegetation diversity [28], we hypothesized that small mammal community metrics would be higher in restored wetlands than in natural wetlands.

2. Methods

2.1. Study Area

We sampled 26 wetlands (14 restored and 12 natural wetlands) across West Virginia, USA, and its three main ecoregions: Ridge and Valley (5 restored and 5 natural), Central Appalachians (4 restored and 2 natural), and Western Allegheny Plateau (5 restored and 5 natural) (Figure 1) [33]; 10 were sampled in 2020 (6 restored, 4 natural) and 16 in 2021 (8 restored, 8 natural). Broad characterizations of these ecoregions are that the Ridge and Valley have forested ridges with agricultural valleys, the Central Appalachians have a higher elevation and increased rainfall, and the Western Allegheny Plateau has many hills and agriculture [34]. Mean annual precipitation for the Ridge and Valley, Central Appalachians, and Western Allegheny Plateau ecoregions are 1138 mm, 1180 mm, and 1063 mm, respectively [35]. Elevation reaches its highest in the Central Appalachians (1402 m), with heights in Ridge and Valley ranging from 152 to 1311 m, and lower elevation in the Western Allegheny Plateau (<610 m) [34].
West Virginia is primarily forested (79%) [36]. Wetlands cover <1% of West Virginia’s land surface, which totals about 40,468 ha [37]. Restored wetlands ranged from 2 to 28.7 ha (mean ± SE ha = 8.1 ± 1.9 ha) and natural wetlands from 1.5 to 45 ha (mean ± SE ha = 12.6 ± 3.6 ha). Wetlands were owned by the West Virginia Division of Natural Resources (4 restored, 8 natural), West Virginia Division of Highways (6 restored, 0 natural), U.S. Forest Service (1 restored, 3 natural), private landowners (2 restored, 1 natural), and the Potomac Valley Audubon Society (1 restored, 0 natural).
We classified wetlands as palustrine emergent (7 restored, 3 natural), scrub-shrub (6 restored, 4 natural), or forested (1 restored, 5 natural) [38]. However, many wetlands had patches of emergent, scrub-shrub, and forested areas. The soil at sampled wetlands was primarily silt loam [39], and all sites had underlying sedimentary geology of shale, sandstone, or alluvium [40]. Elevation ranged from 146 to 660 m (mean ± SE m = 426.5 ± 48.2 m) for restored wetlands and from 167 to 695 m (mean ± SE m = 432.9 ± 57.3 m) for natural wetlands. Restored wetland sites were of differing ages (mean ± SE years = 14.2 ± 2.8 years), with older sites made in 1992 (29 years old) to newer sites restored in 2020 (1 year old).

2.2. Small Mammal Trapping

We used a transect trapping design to capture terrestrial small mammals because they have higher rates of capture, are more efficient at sampling communities [41], and are more likely to capture rare species than a traditional grid array [42]. Our design consisted of 240 m long transects, with traps spaced 10 m apart [43]; therefore, there were 25 traps per transect. We placed transects on both edge and interior locations of larger wetlands (>240 m in length). We placed transects ≥50 m apart and established 2–6 (mean = 3.3; SE = 0.2) transects per wetland. We determined wetland boundaries before trapping by referring to the National Wetlands Inventory geographic information systems (GIS) layer [44] and on-site evaluation of wetland boundaries (using wetland plant and hydrology indicators). Between wetlands, we cleaned equipment to avoid the spread of invasive species and diseases [45].
We used folding Sherman Live Traps (5.08 cm × 6.35 cm × 16.51 cm; H.B. Sherman Traps, Inc., Tallahassee, FL, USA) baited with peanut butter and oats wrapped in wax paper [46]. We replaced bait as needed throughout the trapping session. To encourage small mammal survival, we added cotton to each trap [47]. We checked traps every ≤24 h in the morning during the trapping session. A trapping session consisted of five consecutive nights of trapping. Trapping was restricted to June through August of both sampling years because seasonal and temporal variation among sites can lead to incorrectly drawn conclusions about the small mammal community [48]. To limit the effects of weather and precipitation variation between wetland types, we trapped restored and natural wetlands simultaneously in pairs and in close enough proximity to each other to experience similar weather.
We marked most small mammals with a #1005-1 Monel ear tag (National Band and Tag Company, Newport, KY, USA) in their left ear. We did not ear tag shrews due to the tendency of the tag to quickly fall off and damage their ears [49]. Instead, we marked shrews with hair dye in a unique pattern of dots to distinguish among captured individuals [49]. If a small mammal occupied a trap, we recorded species, mass, body and tail length, sex (male, female), and reproductive condition (adult, juvenile) [50,51]. Additionally, we recorded if females were pregnant or lactating. We differentiated white-footed mice (Peromyscus leucopus) and deer mice (Peromyscus maniculatus) by the body and tail lengths and tail pelage [52].

2.3. Statistical Analysis

2.3.1. Apparent Abundance

We calculated apparent abundance using count data of unique individuals for each observed species and a total count of small mammals across all species. We call this apparent abundance because it is not a direct estimate of abundance but instead assumes the number of individuals captured (counts) is proportional to the actual quantity. We estimated apparent abundance instead of abundance using our capture–mark–recapture data because there were not enough captures to estimate abundance for each species reliably. We evaluated differences in apparent abundance between natural and restored wetlands using a generalized linear model with a Poisson distribution because count data were our response variable. We specified wetland type (restored or natural) as the predictor variable and included an offset for trap nights to standardize trapping effort across sites. To calculate trapping effort (number of trap nights) at each location, we multiplied the number of traps in a transect (25) times the number of transects in a wetland (2 to 6 depending on the site) and multiplied by the number of nights in a trapping session (5). Therefore, each wetland met a minimum of 250 planned trap nights. However, we subtracted a half-trap night from the trap night total when a trap had been falsely snapped and was empty [53,54]. This half-trap night subtraction assumes the trap was open for at least half the night before it closed, leaving the opportunity for a small mammal to enter earlier [53,54]. Using apparent abundance models, we calculated the difference in log expected abundance for each species between restored and natural wetlands. Our hypothesis test included a type 1 error rate of 0.05.

2.3.2. Mass

To determine if the mass of each species differed between wetland types, we compared mass data collected on individuals (initial capture only) by species between natural and restored wetlands with general linear models. Our response variable was mass, and our predictor variables were wetland type, sex, reproductive condition, wetland type by sex interaction, and wetland type by reproductive state interaction. We included interaction terms to conclude whether adult, juvenile, female, or male mass differed by wetland type. We used contrasts to account for the influence of sex (male and female) and reproductive condition (juvenile and adult) when comparing mass across wetland types. We calculated contrasts using the package ‘multcomp’ in program R [55,56].

2.3.3. Occupancy

Given low capture rates, we compared the occupancy probability of small mammals between wetland types. We ran single-season occupancy models since we only visited each site once for a 5-night trapping session [57]. Our response variable was the detection or non-detection of a species at a site. We treated each transect as a replicate survey, having 2–6 replicate surveys per site. We collapsed species detection within a transect as a 1 if detected and a 0 if not detected over the 5-night trapping session per site. Wetland type, restored or natural, was used as our site-level covariate, and our survey level covariate (detection covariate) was held constant. We implemented these models using a maximum likelihood estimation approach with the R package “unmarked” [58]. We fit occupancy models for all taxa except deer mice; we detected them at each site, which precludes the use of occupancy analysis.

2.3.4. Community Composition

To assess differences in small mammal community composition, we used non-metric multidimensional scaling (NMDS). This technique uses rank orders, making it more flexible and accommodating more data types than other ordination methods [59]. We specified Bray–Curtis dissimilarity to obtain a distance matrix which we then plotted for a visual assessment of community composition between the two wetland types. To numerically assess differences in community composition between wetland types, we used an analysis of similarity (α = 0.05) to complement the NMDS plot [60,61] and again specified Bray distance. Community data for the analysis was our raw count data for each species for each site, with each value standardized by catch-per-unit effort (CPUE). We calculated this as the percentage of captured individuals divided by the trapping effort at a given site (i.e., read as the number of captures/100 trap nights) [62]. To implement NMDS, we used the function metaMDS in vegan [63] and specified two reduced dimensions with 100 iterations until a solution was reached. We then used the function anosim in vegan [63] to analyze small mammal similarity.

3. Results

After adjusting for snapped traps, we had a total trapping effort of 10,060 trap nights across our 26 wetland sites over 2 summers. Trapping effort ranged from 204.5 to 719 trap nights (mean ± SE = 412.8 ± 40.7) at restored wetlands and 214 to 584 trap nights (mean ± SE = 356.6 ± 42.57) at natural wetlands. There were 6.36 captures per 100 trap nights (640 total captures; 349 in natural and 291 in restored wetlands), and 4.25 captures per 100 trap nights were unique individuals (428 unique individuals; 218 in natural and 210 in restored wetlands). Recaptured individuals comprised 2.10 captures per 100 trap nights (212 recaptures; 131 in natural and 81 in restored wetlands) and were primarily Peromyscus spp. Nine total species were represented. We captured deer mice (N = 187), white-footed mice (N = 111), meadow voles (N = 73), meadow jumping mice (N = 16), northern short-tailed shrews (N = 23), and eastern chipmunks (N = 11) at both wetland types, and woodland jumping mice (Napaeozapus insignis) (N = 2), masked shrews (Sorex cinereus) (N =4), and a southern flying squirrel (Glaucomys volans) (N =1) exclusively at natural wetlands.

3.1. Apparent Abundance

Peromyscus was the most captured genus, with deer mice accounting for 43.69% of total unique captures and white-footed mice 25.93%. Apparent abundance of deer mice was greater for natural wetlands (Z = 4.84, p < 0.001) (Table 1), but similar between wetland types for white-footed mice (Z = 1.488, p = 0.137). Meadow voles accounted for 17.05% of unique individuals captured. The apparent abundance of meadow voles was greater in restored wetlands, but evidence of a difference between wetland types was marginal (Z = −1.89, p = 0.06). We captured northern short-tailed shrews, meadow jumping mice, and eastern chipmunks less frequently (5.37%, 3.73%, and 2.57% of unique captures, respectively), and we did not see a difference in apparent abundance between wetland types (p = 0.246, 0.684, and 0.170, respectively). Woodland jumping mice, masked shrews, and the southern flying squirrel were captured only in natural wetlands and were our three least common captures (0.46%, 0.93%, 0.23%, respectively).

3.2. Mass

We found no differences in the mass of deer mice, white-footed mice, meadow voles, northern short-tailed shrews, eastern chipmunks, and meadow jumping mice between restored and natural wetlands when compared by age-sex cohort (Table 2). Numerically, mean body mass tended to be higher for deer mice, white-footed mice, and meadow jumping mice in restored wetlands and higher for meadow voles, short-tailed shrews, and chipmunks in natural wetlands; however, none of the differences were statistically different from zero (Table 2).

3.3. Occupancy

Occupancy probability was similar between wetland types for white-footed mice (Z = 0.315, p = 0.753), meadow voles (Z = −0.287, p = 0.774), northern short-tailed shrews (Z = 0.119, p = 0.905), meadow jumping mice (Z = −0.018, p = 0.985), and eastern chipmunks (Z = 0.216, p = 0.828) (Table 3).

3.4. Community Composition

NMDS analysis showed substantial overlap in small mammal community composition between restored and natural wetlands (Figure 2). The stress value associated with the distance matrix used to make the plot for small mammal community was 0.19. This value is regarded as fair but still valid because it signifies the potential for the analysis to be misleading; traditionally, stress values are most reliable when they are <0.05 [64]. However, increasing the dimensions beyond two did not significantly reduce stress or alter the results. The similarity analysis indicated that small mammal community composition was similar between wetland types (R = 0.032, p = 0.226).

4. Discussion

We hypothesized that small mammal community metrics would be higher in restored wetlands than in natural wetlands. However, our results did not support our hypothesis. Restored wetlands contained similar terrestrial small mammal communities as natural wetlands, though not equivalent. Two main discrepancies between wetland types were discovered, suggesting natural wetlands were more favorable to the small mammal community. One difference was that natural wetlands supported a greater apparent abundance of deer mice. Another difference was that only six species were found in restored wetlands, as opposed to nine species found in natural wetlands, suggesting natural wetlands may have a greater ability to host a wider variety of terrestrial small mammal species. In terms of small mammal communities, restored wetlands do appear to be successful in providing adequate habitat for terrestrial small mammals, though they are certainly not equivalent to natural wetlands.

4.1. Apparent Abundance, Occupancy, and Mass

We found apparent abundance of deer mice was higher in natural wetlands. The average expected count for deer mice per 100 trap nights for natural wetlands was more than double the estimate for restored wetlands. We possibly observed a higher apparent abundance of deer mice in natural wetlands because natural wetlands may provide more desirable habitat for small mammals, as evidenced by the three species we found in natural wetlands but not restored wetlands. Excluding deer mice, our results are consistent with Kurz et al. [31], who also found no difference in relative abundance of small mammals between restored and reference wetlands (note that Kurz et al. [31] did not include deer mice in their analysis).
Meadow voles were bordering a significant difference in apparent abundance between wetland types; therefore, this result may be of biological significance and would likely be significant with a larger sample size, specifically in younger restored wetlands. The apparent abundance of the Meadow vole was 61% higher in restored wetlands than in natural wetlands. Although meadow voles are a common non-obligate wetland species [65] and were the most abundant small mammal species in another West Virginia wetland [66], our nearly significant finding may be related to wetland age. Meadow voles have a higher apparent abundance at younger wetlands [67], presumably because younger wetland sites were typically palustrine emergent which better corresponds to their habitat affinity for grassland, as opposed to woody vegetation [68,69]. Specifically, younger wetlands (< 5 years old) had more grasses and less woody vegetation, while older sites (> 10 years old) were characterized by more woody vegetation because they had more time to become established [67]. However, our hypothesis that restored wetlands supported higher vegetative species diversity was not sustained, and results indicated no differences in vegetation species diversity or community composition between natural and restored wetlands [67].
Unlike deer mice, we could not determine a difference in apparent abundance between wetland types for white-footed mice, the other Peromyscus species captured. Given both species are similar in that they share the same food resources and exist in similar microhabitats [70], it was unexpected that they did not have a higher average apparent abundance in natural wetlands like that of deer mice. Variations in food availability and weather conditions could change the competitive advantage; deer mice are efficient at using torpor during food shortages which increases their survival in years of poor food production compared to white-footed mice [71]. This variability potentially leads to one species being more common than the other and may explain why we encountered more deer mice than white-footed mice. By finding a similar apparent abundance of white-footed mice, we may assume that vegetative structural complexity is similar between both restored and natural wetlands, as structural complexity is important to this species [72].
Northern short-tailed shrews are prevalent in this region and are not specific to wetlands or other habitat types [67,73]. Therefore, it is unsurprising that we did not find a difference between wetland types. Meadow jumping mice are more habitat-specific in that they prefer moist areas [74,75]. However, wetland types had no apparent abundance or occupancy probability differences. Although occupancy was similar between wetland types for northern short-tailed shrews and meadow jumping mice, detection probability was <0.3, creating more uncertainty in the results for these species. Meadow jumping mice associate with jewelweed (Impatiens capensis) [76], which we frequently observed in both wetland types. Both northern short-tailed shrews and meadow jumping mice have no clear preference in terms of the vegetation type chosen as shelter (herbaceous, shrubby, or woody vegetation) [77]; therefore, any differences in vegetation type of restored wetlands in comparison to natural wetlands is not likely to affect the wetland’s ability to provide shelter for these species.
Ecologists commonly use mass to evaluate the environment of small mammals [78,79]. We expected differences in small mammal mass if food abundance, such as insect community or mast production, differed between wetland types. Because we found mass to be similar between wetland types, this suggests that the ecological condition of wetland types is similar. Although, small mammal mass may be insensitive to environmental perturbations, as it is unaffected by patch size or pollution [80,81].

4.2. Community Composition

We found that the terrestrial small mammal community composition was similar between wetland types. This finding was encouraging because policies such as no net loss are only concerned with totals and overlook species composition changes [82]. Like our findings, Kurz et al. [31] also did not discover differences in community composition when researching small mammal communities between restored and natural wetlands. Sites that did not overlap in terms of terrestrial small mammal community had no clear commonalities with each other, aside from all being natural wetlands; these non-overlapping wetlands were not specific to the ecoregion, sampling year, or dominant vegetation type.
Although community composition overlapped between the wetland types, we also observed species variations. Three of the nine species we caught were found only in natural wetlands: 4 masked shrews, 2 woodland jumping mice, and 1 southern flying squirrel encountered throughout the study. It is noteworthy, though, that these were our three rarest species in terms of capture, which is likely why our community composition analysis did not find a difference between wetland types. We may not have captured masked shrews and woodland jumping in restored wetlands because they are sensitive to habitat modification; Racey and Euler [83] found both species were sensitive to development. Potentially, both species left the wetland area when it was a degraded wetland in the years before restoration, and they simply have yet to re-colonize the patch. In addition to being a sensitive species [83], the woodland jumping mouse has strong associations with a high volume of decomposed logs [84]. These decomposed logs are also where fungi, such as Endogone spp., Melanogaster spp., and Hymenogaster spp., an important food source for woodland jumping mice, grow [84,85,86]. Conditions for this fungus were perhaps more favorable in natural wetlands, where trees had more time to grow and decompose than in restored wetlands. We captured one southern flying squirrel only once, and it was at a natural wetland. Perhaps we caught it only in a natural swamp because the species prefers mature forests [87]. The oldest restored wetland sampled is approaching 30 years old, which may be too young still to have a mature forest.
Most species we captured were habitat generalists, like the findings of Francl et al. [88] in high-elevation wetlands. However, we did catch a habitat specialist, woodland jumping mice [84]. Because we only captured them in natural wetlands, it is still unclear whether restored wetlands can support habitat specialists. However, woodland jumping mice are not wetland-obligate species or wetland specialists but are a specialist in forests. Similarly, Francl et al. [88] found that captured habitat specialists were those from a surrounding habitat type in high elevation wetlands.
Most of our captured species are secure in their populations within the state and globally [89]. All species except for meadow jumping mice and woodland jumping mice are in the secure category (ranked S5) [89]. Meadow jumping mice, whose populations are vulnerable in the state (ranked as S3) [89], were found in both wetland types and were similar in apparent abundance and occupancy between wetland types. Woodland jumping mice, whose populations are uncommon but not rare in the state (ranked S4) [89], were only captured twice in the study, and both were in natural wetlands. Although meadow jumping mice (S3 species) can be supported by restored wetlands similarly to natural wetlands, only natural wetlands supported woodland jumping mice (an S4 species).

5. Conclusions

We conclude that restored wetlands can be an adequate treatment for replacing natural wetlands in terms of small mammal communities but are not equivalent to natural wetlands. We did not discover differences between wetland type for mass, occupancy probability, or community composition. These results are encouraging because small mammals are traditionally not considered in wetland restoration efforts. However, we did find that natural wetlands supported a higher apparent abundance of deer mice. We also discovered our three most rarely captured species were found exclusively in natural wetlands and did not find wetland-obligate species in restored wetlands. Thus, future research should focus on wetland-obligates and rare species in restored wetlands to discern whether they can provide quality habitats for these species. Reconsidering compensation types and ratios for government-mandated compensatory wetland mitigation programs in the United States and other countries may help to bridge this gap. Regulations requiring more wetland replacement acreage per acre lost or requiring the same wetland type to be replaced as the lost wetland may provide more habitat for less common species. We conclude that restored wetlands can mostly provide for small mammal communities like natural wetlands but are not equivalent to natural wetlands.

Author Contributions

Conceptualization, J.T.A. and K.L.N.; methodology, K.L.N., J.T.A., C.T.R. and M.W.F.; formal analysis, K.L.N. and C.T.R.; investigation, K.L.N. and J.T.A.; data curation, K.L.N.; writing—original draft preparation, K.L.N.; writing—review and editing, J.T.A., C.T.R. and M.W.F.; visualization, J.T.A. and K.L.N.; supervision, J.T.A.; funding acquisition, J.T.A. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the National Science Foundation, grant number 01-1458952A; the USDA National Institute of Food and Agriculture McStennis Project, grant number WVA00117; WVU Natural History Museum, West Virginia Division of Natural Resources, the West Virginia Division of Highways, and the James C. Kennedy Waterfowl and Wetlands Conservation Center at Clemson University.

Institutional Review Board Statement

This study was approved by the Institutional Animal Care and Use Committee Review Board of West Virginia University (Protocol #2004034461).

Data Availability Statement

The data used in this study can be obtained from the authors upon request.

Acknowledgments

We thank the public and private landowners for granting access to the wetland sites used in this study. Additionally, we thank Joseph Groome for his help in the field. We thank Dave Deane and an anonymous reviewer for their helpful comments.

Conflicts of Interest

The authors declare no conflict of interest.

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Land 11 01482 g001 550
Figure 1. Twenty-six wetland sites (14 restored and 12 natural) were sampled in 2020 and 2021 for small mammal and vegetation communities throughout West Virginia, USA. Sites are plotted against the state’s three ecoregions: Ridge and Valley, Central Appalachians, and the Western Allegheny Plateau.
Figure 1. Twenty-six wetland sites (14 restored and 12 natural) were sampled in 2020 and 2021 for small mammal and vegetation communities throughout West Virginia, USA. Sites are plotted against the state’s three ecoregions: Ridge and Valley, Central Appalachians, and the Western Allegheny Plateau.
Land 11 01482 g001
Land 11 01482 g002 550
Figure 2. Non-metric multidimensional scaling (NMDS) scores for small mammal community composition from restored (n = 14) and natural (n = 12) wetland sites in West Virginia, USA, sampled from 2020–2021. Convex hulls surround restored wetland communities (blue) and natural wetland communities (pink). We specified 2 dimensions with 100 iterations. The distance matrix stress value was 0.19. Analysis of similarity indicates no significant difference in small mammal communities by wetland type (R = 0.032, p = 0.226).
Figure 2. Non-metric multidimensional scaling (NMDS) scores for small mammal community composition from restored (n = 14) and natural (n = 12) wetland sites in West Virginia, USA, sampled from 2020–2021. Convex hulls surround restored wetland communities (blue) and natural wetland communities (pink). We specified 2 dimensions with 100 iterations. The distance matrix stress value was 0.19. Analysis of similarity indicates no significant difference in small mammal communities by wetland type (R = 0.032, p = 0.226).
Land 11 01482 g002
Table
Table 1. Mean and standard error of the apparent abundance of unique small mammals per 100 trap nights by wetland type sampled in the summers of 2020 and 2021 in West Virginia, USA. Over both years, 3 species (woodland jumping mouse, masked shrew, and southern flying squirrel) were only observed in natural wetlands and were excluded from the table.
Table 1. Mean and standard error of the apparent abundance of unique small mammals per 100 trap nights by wetland type sampled in the summers of 2020 and 2021 in West Virginia, USA. Over both years, 3 species (woodland jumping mouse, masked shrew, and southern flying squirrel) were only observed in natural wetlands and were excluded from the table.
Restored (n = 14)Natural (n = 12)
SpeciesMeanSEMeanSE
Peromyscus maniculatus1.2801.1232.6401.098
Peromyscus leucopus0.9681.1421.2851.144
Microtus pennsylvanicus0.8651.1510.5371.231
Blarina brevicauda0.2761.2840.1631.459
Zapus hudsonius0.1731.3710.1400.150
Tamias striatus0.0691.6480.1631.459
Table
Table 2. Mass model contrast results for each small mammal species captured in both wetland types (n = 6), for adult males (AM) and females (AF), and juvenile males (JM) and females (JF). Small mammals were captured in West Virginia, USA, in the summers of 2020 and 2021. Species with missing results for categories indicate that no captures were made for that category to compare.
Table 2. Mass model contrast results for each small mammal species captured in both wetland types (n = 6), for adult males (AM) and females (AF), and juvenile males (JM) and females (JF). Small mammals were captured in West Virginia, USA, in the summers of 2020 and 2021. Species with missing results for categories indicate that no captures were made for that category to compare.
Peromyscus maniculatusPeromyscus leucopusMicrotus pennsylvanicusBlarina brevicaudaTamias striatusZapus hudsonius
Age-Sex Cohortt-Valuep-Valuet-Valuep-Valuet-Valuep-Valuet-Valuep-Valuet-Valuep-Valuet-Valuep-Value
AM−0.2130.9961.2370.551−0.0671.000−0.3260.935−0.0970.994−0.7890.456
AF0.0871.0001.4670.404−0.7420.855−0.4740.868−0.5160.848--
JM0.5410.936−0.1490.999−0.5670.928------
JF0.5870.9200.3930.974−0.8510.798------
Table
Table 3. Occupancy probabilities by wetland type and detection probabilities for five species captured in restored (n = 14) and natural (n = 12) wetlands throughout West Virginia, USA, from 2020–2021. Peromyscus maniculatus was omitted from occupancy modeling because it was discovered at each site, leading to poor occupancy estimations.
Table 3. Occupancy probabilities by wetland type and detection probabilities for five species captured in restored (n = 14) and natural (n = 12) wetlands throughout West Virginia, USA, from 2020–2021. Peromyscus maniculatus was omitted from occupancy modeling because it was discovered at each site, leading to poor occupancy estimations.
Occupancy ProbabilityDetection
Probability
RestoredNatural
SpeciesEstimateSEEstimateSEEstimateSE
Peromyscus leucopus0.6510.1540.7240.1800.4630.081
Microtus pennsylvanicus0.5680.1540.5030.1760.5010.087
Blarina brevicauda0.6890.2350.9980.8510.2300.062
Zapus hudsonius0.6330.3930.6250.4300.1620.097
Tamias striatus0.1640.1110.2000.1350.4580.220
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Noe, K.L.; Rota, C.T.; Frantz, M.W.; Anderson, J.T. Restored and Natural Wetland Small Mammal Communities in West Virginia, USA. Land 2022, 11, 1482. https://0-doi-org.brum.beds.ac.uk/10.3390/land11091482

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Noe KL, Rota CT, Frantz MW, Anderson JT. Restored and Natural Wetland Small Mammal Communities in West Virginia, USA. Land. 2022; 11(9):1482. https://0-doi-org.brum.beds.ac.uk/10.3390/land11091482

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Noe, Krista L., Christopher T. Rota, Mack W. Frantz, and James T. Anderson. 2022. "Restored and Natural Wetland Small Mammal Communities in West Virginia, USA" Land 11, no. 9: 1482. https://0-doi-org.brum.beds.ac.uk/10.3390/land11091482

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