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All issues Volume 58 (2022) Int. J. Lim., 58 (2022) 16 Full HTML
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Int. J. Lim.
Volume 58, 2022
Article Number 16
Number of page(s) 10
DOI https://doi.org/10.1051/limn/2022016
Published online 23 December 2022

© EDP Sciences, 2022

1 Introduction

One of global challenges is the conservation of freshwater ecosystems, where biodiversity is declining much faster than in marine and terrestrial ecosystems (WWF, 2012). Major threats to freshwater ecosystems include overexploitation (removal of living resources), water pollution, habitat degradation, and invasions of non-native species, all of which are interconnected (Dudgeon et al., 2006). Among freshwater ecosystems, small water bodies need special protection because they play a disproportionally large role in global geochemical cycles (Downing, 2010).

Some of the most widespread small water bodies are floodplain lakes, which provide a variety of habitats for many groups of organisms such as invertebrates, fish, amphibians, and birds (Joniak and Kuczýnska-Kippen, 2016). Floodplain water bodies support the structure of phyto- and zooplankton communities in all water objects of a river system, including the main watercourse (Krylov, 2015). Such water bodies are important for the conservation and reproduction of fish because floodplains are spawning and nursery grounds for many fish species and serve as refugia for them during floods (Naus and Adams, 2018).

The current global climate processes pose many threats that lead to changes in temperature and water flow, thereby causing eutrophication and alteration of food chains (Golubkov, 2021). In recent decades, intra-annual redistribution of seasonal runoff (Blöschl et al., 2019) and a reduction in spring flood volumes (Frolova et al., 2015) became usual for European river valleys. These factors, together with anthropogenic disturbance of the hydrologic regime of rivers, lead to the cessation of water exchange between floodplain water bodies and the main river and among themselves; this problem could result in significant changes in aquatic communities. Because of specific morphological and abiotic properties, floodplain lakes are highly susceptible to degradation, and the shallowing of these ecosystems may be accelerated by human activities (Koc et al., 2009). Low frequency of high floods can result in a decrease in lateral connectivity between floodplain lakes and the river, thereby possibly affecting fish biodiversity negatively (Glińska-Lewczuk et al., 2016). The loss of water bodies variety and heterogeneity can influence aquatic organisms, macrophytes, and periphyton (Hill et al., 2017; Pander et al., 2018). The diversity and mosaicity of habitats within a floodplain contribute to all biodiversity patterns (Amoros and Bornette, 2002).

The influence of the Eurasian beaver (Castor fiber Linnaeus, 1758) and North American beaver (Castor canadensis Kuhl, 1820) on aquatic ecosystems is common in small rivers and streams, where beavers change the ecosystems from lotic to lentic (Rosell et al., 2005; Brazier et al., 2020). The creation of dams on inflows and outflows of lakes also enhances littoral and subsequent effects on abiotic conditions and on aquatic biota (Vehkaoja et al., 2015; Kivinen et al., 2020). Another important aspect of the beaver impact is the digging activity. Canal construction results in an additional open water surface and increases connectivity and heterogeneity (Grudzinski et al., 2019). The creation of burrows may also have large implications for aquatic ecosystems, although the bulk of existing knowledge concerns the use of burrows by other organisms as shelters or dwellings (Dyakov, 1975; Samas, 2015). The impact of beaver burrows on the water body itself remains a poorly investigated issue.

Despite many publications that describe the beaver settlements in lakes, the specificity of beaver activity in lakes is not well studied (Bashinskiy, 2020). The assessment of the beaver influence on lotic ecosystems is usually carried out via a comparison of beaver ponds with an unaffected stream. Such an approach is not suitable for most of lentic ecosystems because it is difficult to choose correct control habitats (e.g., a lake without beavers) without assuming that initial water bodies were identical before beaver settlement. In diverse and dynamic floodplain ecosystems, such assumptions are not correct. Therefore, a potential solution is to compare the habitats within one water body. Although many aquatic organisms can move freely throughout the habitats, their substantial preference for specific microhabitats may indicate more favorable conditions.

The aim of this work was to assess the impact of the beaver digging activity on habitat structure and aquatic organisms in a small floodplain waterbody. To this end, the following tasks were undertaken: assessment of alteration of water body morphometry by the beaver activity, analysis of abiotic characteristics of beaver-created microhabitats, description of plant communities in beaver and control microhabitats, and analysis of the distribution of aquatic organisms in the water body under the influence of zoogenic mosaicity.

2 Material and methods

2.1 Study area

This study was conducted in June 2–8 and July 23–29, 2021, in Penza Oblast, Russia, near Privolzhskaya Lesostep Nature Reserve (52°48′58.4′′ N, 44°27′40.4′′ E) (Fig. 1). The model water body is a part of the old riverbed of the Khoper River (Black Sea basin). This floodplain is under regular monitoring, and high floods are rare. The connection with the main river occurs once in a decade; hence, most water bodies are temporary (Bashinskiy et al., 2019). Beavers have inhabited the water body previously and have dug six burrows along its banks. The last inhabitation of the water body was documented in 2015. Since then, they used the water body only for occasional feeding, without landscape-making activities (unpublished data). During this period, the water level dropped by ∼45% relative to the level documented previously. The entrances to the burrows became exposed, and the ground partially or completely collapsed. These events changed the relief of the shoreline, and therefore new microhabitats appeared in the littoral. These beaver-created sites (BSs) had the shape of small bays characterized by greater depth and steeper underwater slopes. The space between these sites was typical for the unaffected littoral of other floodplain water bodies (control sites: CSs).

thumbnail Fig. 1

The study area and the location (white circle) of the examined water body (Russia, Penza Oblast). The white line denotes the borders of Privolzhskaya Lesostep Nature Reserve.

2.2 Sampling and analytical methods

To understand the scale and status of the beaver activity in the floodplain complex, all water bodies of the old riverbed were examined during the relevant season. Their perimeters were measured with a Garmin GPSmap 60Cx. The status of beaver settlements (active, abandoned, or always absent) was assessed in each water body judging by the presence of fresh cuttings and tracks. All beaver burrows were counted.

A Deeper Smart Sonar CHIRP+ echo sounder, adapted for shallow water, was used to study morphometry of the model water body. The depth data were collected from 567 points (a radial grid was employed) to create a bathymetric map in QGIS 3.18.2 by means of the Contour module (Crook and Roubeyrie, 2017). In addition, the main morphometric characteristics of the water body were calculated: area, the perimeter, the lake basin shape factor (the ratio of average to maximum depth), shoreline development (the ratio of the length of the shoreline to the circumference of the circle having area equal to that of the lake). For determining pre-beaver shoreline, the initial perimeter was calculated, excluding the bays that were formed by collapsed burrows.

Based on the bathymetric map and shoreline morphology, BSs and CSs were identified and measured using QGIS geoprocessing tools. Five BSs and four CSs were analyzed. CSs were situated at ≥2.7 m distance from a BS, the mean distance to the closest BS was 5.1 m. Their area, distances between each other, and the proportion of such habitats in the entire water body were calculated. Besides, four littoral zones were outlined in QGIS at distances of 1, 2, 3, and 3 m from the shore, where the proportion of BSs was assessed too.

At each studied site, water temperature was measured with a digital thermometer. Macrophytes were described using 1 m2 sampling sites. The plant cover (%) and species number were determined within all BSs and CSs and for the entire water body. The guide by Lisitsyna and Papchenkov (2000) was used for the identification of macrophytes.

Funnel traps were utilized to investigate aquatic organisms because these traps are the most universal (Skelly and Richardson, 2010) and result in little habitat disturbance. To improve the effectiveness of catching, two different traps were used. One (5 mm mesh, 700 mm length, and an inlet diameter of 120 mm) was more suitable for adult fish and amphibians and large tadpoles. The other (2 mm mesh, 350 mm length, and an inlet diameter of 60 mm) was aimed at catching macroinvertebrates and small tadpoles. At each point, the funnel traps stayed for 3 days. Because burrow BS5 dried up in July, the traps were moved to burrow BS4. Forty-eight captures were made during 12 days in the study period. We identified species (genera for insects) by means of a guide to zooplankton and zoobenthos of freshwaters of European Russia (Alekseev and Tsalolikhin, 2016). Amphibian larvae were weighed, and their stage of development was determined according to Gosner (1960). All caught organisms were released back into the water in the opposite part of the water body, where there were no traps. For analysis of relative abundance, the number per catching effort was used, meaning all caught individuals per day.

2.3 Statistical analyses

These procedures were performed in STATISTICA 7.0 software (StatSoft, 2004) and Past 3.16 software (Hammer et al., 2001). The nonparametric Spearman rank correlation (RS) coefficient was used for correlation analysis (Myers et al., 2010); the nonparametric Mann–Whitney U test was employed for evaluating differences between samples (Mann and Whitney, 1947); and canonical correspondence analysis for multivariate analysis (Legendre and Legendre, 1998). The preference of a species for different habitat types was assessed by the chi-square test (Sokal and Rohlf, 1981). For identifying species indicative of given groups of habitats, we performed the indicator species analysis (IndVal) (Dufrêne and Legendre, 1997). The Jaccard (J) index was used to measure the similarity of species diversity between different habitats (Magurran, 1988).

3 Results

The main morphometric characteristics of the water body in question are given in Table 1, and the bathymetric map in Figure 2. The total area of BSs reached 62.2 m2, which was 7% of the area of the whole water body. They contributed to 30% of all littoral habitats at 1 m distance from the shore, to 29% at 2 m distance, 26% at 3 m distance, and 23% at 4 m distance. In addition, the beaver activity increased the perimeter of the water body by 8 m (10%) relative to the initial perimeter. Furthermore, shoreline development increased from 0.98 to 0.91 relative to the pre-beaver shoreline.

The significant difference between BSs and CSs was greater depth in June (U test, p < 0.001). The difference was not significant in July, when the microhabitats became more similar because of gradual drying out (Fig. 3a). Despite the different depths, water temperatures were almost the same within a month (Fig. 3b), but there was a temperature difference between months (U test, p < 0.001).

The second important difference between habitats was plant cover (Fig. 3c), which differed significantly between BSs and CSs (U test, p < 0.001). The situation changed by July, and the difference decreased but was still significant (U test, p = 0.034). This pattern was also characteristic of species diversity of flora. Ten species of vascular plants were found within the boundaries of the water body, six of which grew in the studied microhabitats (Appendix A). In addition, in June, the water body was overgrown by filamentous algae of the genus Spirogyra. Although the total plant cover of all sites remained almost unchanged, in June, all examined microhabitats had vegetation, whereas in July, three sampling sites were free of macrophytes. The average number of plant species diminished in the littoral, primarily because of a decrease in emergent vegetation diversity of CSs (Fig. 4). In BSs, the total number of species remained roughly the same, although the variance of values increased. In June, emergent vegetation was represented mainly by one species (Alisma plantago-aquatica) in BSs, whereas in July, its diversity went up in these habitats. The depth and size of BSs negatively correlated with the number of macrophyte species (RS = −0.29, p = 0.042, and RS = −0.42, p = 0.003, respectively). Depth and size determined the distribution of Alopecurus aequalis (RS = −0.63, p < 0.001 for depth, RS = −0.48, p < 0.001), which significantly preferred CSs (χ2 = 20.2; df =1; p < 0.001). The same was true for the other widespread helophyte, Sparganium erectum, for which beaver habitats were less favorable (χ2 = 4.0; df =1; p = 0.046). The opposite was true for Potamogeton lucens, which was found to be confined to beaver microhabitats in the littoral zone (χ2 = 6.86; df =1; p = 0.009). The indicator species analysis confirmed preferences of some emergent plants for unaffected habitats in June: Sparganium erectum (IndVal = 56.3%, p = 0.024) and Alopecurus aequalis (IndVal = 80%, p = 0.003).

Twenty-four taxa of aquatic organisms were detected according to the results of capture by the funnel traps (Appendix B). The taxon lists of the two types of microhabitats largely overlapped, and most of invertebrate species used different microhabitats equally (Figs. 5a, Figs. 5c). The key factors influencing the distribution of organisms were the depth and plant cover of the different sites. More insect species were observed in BSs (RS = 0.34, p = 0.022) for both months. The depth and size of BSs were important determinants of this parameter. Notonecta sp. and Naucoris sp. were more abundant at deeper sites (RS = 0.34, p = 0.016, and RS = 0.34, p = 0.019, respectively). Relative abundance of beetles Dytiscus sp. and Hydrophilus sp. (RS = 0.29, p = 0.043, and RS = 0.27, p = 0.046, respectively) correlated with sizes of BSs. Naucoris sp. and Hydrophilus sp., preferred BSs statistically significantly in June (χ2 = 4.8; df =1; p = 0.029, for both taxa). Mollusks Planorbis planorbis (RS = 0.28, p = 0.041) and Lymnaea palustris (RS = 0.31, p = 0.050) and beetles Hydrous sp. (RS = 0.30, p = 0.038) were found to prefer thickets of emergent vegetation: Alopecurus aequalis, Alisma plantago-aquatica, and S. erectum, respectively.

The correspondence analysis of fish and amphibians also revealed an overlap between taxon lists of the two microhabitat types (Figs. 5b, Figs. 5d). In contrast to macroinvertebrates, depth played a lesser role, and BS size was more important. Nonetheless, some species displayed significant preferences. In June, adult crucian carps Carassius carassius preferred CSs (χ2 = 4.8; df =1; p = 0.029). Although in July, the abundance of adults of that fish was also high in BSs, the difference from CSs was not significant. Adult weatherfishes Misgurnus fossilis were found only in BSs in June (IndVal = 50%, p = 0.048), but their juveniles were found only in CSs in July (IndVal = 50%, p = 0.045). Juvenile crucian carp avoided thickets of emergent plants Alisma plantago-aquatica (RS = −0.67, p < 0.001) and Alopecurus aequalis (RS = −0.45, p = 0.003) preferring submerged vegetation P. lucens (RS = 0.35, p = 0.015).

Adult marsh frogs Pelophylax ridibundus occurred mostly in BSs (χ2 = 6.2; df =1; p = 0.013) in June. In July, frog tadpoles avoided BSs (χ2 = 4.2; df =1; p = 0.041) and preferred CSs (IndVal = 53.6%, p = 0.016). One newt Lissotriton vulgaris was found in a BS. Adult fire-bellied toads Bombina bombina preferred CSs (IndVal = 50%, p = 0.048). No significant difference between CSs and BSs in any parameters was found for tadpoles of the spadefoot Pelobates vespertinus. In June, on average, more tadpoles preferred CSs, but their biomass was higher near BSs by 0.3 g on average (Fig. 6). By July, all parameters equalized, and the abundance of tadpoles in BSs rose (IndVal = 45.2%, p = 0.034), but they lagged in terms of Gosner's stages of development by one stage on average (Fig. 6). All three parameters correlated with water temperature: RS = 0.68, p = 0.006 (abundance), RS = 0.71, p = 0.004 (biomass), and RS = 0.58, p = 0.025 (stage of development). As noted above, temperature did not depend on beaver activity.

In addition, overall species similarity between different habitats was analyzed. Distances between sites did not influence the similarity of habitats (RS = 0.14, p = 0.757, in June; RS = −0.31, p = 0.462, in July). Overall, microhabitats differed (J = 0.48), with BSs being the most similar among themselves (J = 0.56). CSs differed more among themselves (J = 0.52) and from BSs (J = 0.49). The beaver activity slightly raised β-diversity within the water body: Jaccard indices changed from 0.48 to 0.52 after the appearance of zoogenic microhabitats.

Table 1

The main morphometric characteristics of the studied water body in June–July.

thumbnail Fig. 2

The bathymetric map of the water body in question (June, 2021). The dashed line represents the pre-beaver shoreline (initial perimeter). The hatched areas show beaver-created microhabitats (BS1–BS4). CS1–CS5 are control sites. The red lines indicate littoral zones at distances of 1, 2, 3, and 4 m from the shore.
thumbnail Fig. 3

Main characteristics of microhabitats of different types (white boxes: control, gray boxes: created by beavers). Depth (a), water temperature (b), and plant cover (c).
thumbnail Fig. 4

Changes in the number of all plant species (a) and of emergent (b) and submerged plants (c) in microhabitats. White boxes are CSs; gray boxes are BSs.
thumbnail Fig. 5

A species-conditional triplot based on the canonical correspondence analysis for invertebrate (a: June, c: July) and vertebrate (b: June, d: July) biota. A: adults and imago individuals, J and L: larvae and juvenile individuals. Black dots show BSs, and gray dots represent CSs. SIZE: the size of microhabitat altered by beavers, TEMP: water temperature, COVER: plant cover.
thumbnail Fig. 6

Relative abundance (a), biomass (b), and the stage of development (Gosner, 1960) (c) of spadefoot tadpoles.

4 Discussion

In floodplain water bodies, beaver digging may be of key importance (Bashinskiy, 2020). Creation of burrows and channels by beavers improves habitat heterogeneity and connectivity (Hood and Larson, 2014). Beaver digging supports the mosaic of microhabitats, thereby creating spatial heterogeneity, which is one of the most important properties of ecosystems and can determine their functioning (Turner and Chapin, 2005). Many research articles indicate that small-scale spatial structure is one of the most important determinants of the composition of invertebrate communities (Kuczyńska-Kippen and Nagengast, 2006; Burgazzi et al., 2020) and amphibian communities (Sun et al., 2021). Our paper shows that beavers improve heterogeneity of littoral habitats and improve β-diversity.

The hydroperiod plays an important role in the metamorphosis of amphibiotic insects and amphibians (Pechmann et al., 1989; Brooks, 2000; Schriever et al., 2014). A shortening of the period of a water body's presence could lead to the death of the larvae. Although our study does not show the importance of digging for tadpole survival, this zoogenic factor needs to be considered in the face of modern degradation of floodplain ecosystems, the shortening of the hydroperiod, and transformation of permanent water bodies into temporary ones. Some authors have documented the deepening of water bodies by beavers (Pankov and Pankova, 2016; Nitsche, 2021); sometimes, their digging creates new ponds (Westbrook et al., 2017).

Our results point to a significant decrease in the diversity of emergent plants near beaver-altered microhabitats, mainly because of their depth. Overgrowth of water bodies is usually provoked by such vegetation in the littoral (e.g., Skowron and Jaworski, 2017; Lawniczak-Malińska et al., 2018). Thus, the observed beaver-driven ∼30% reduction of habitats suitable for emergent plants may be an obstacle to the shallowing of the water body. Previously, Pankova and Pankov (2018) noted that shoreline vegetation is less common in locations with beaver burrows than in locations not suitable for digging in oxbows. By contrast, the deeper locations of our studied water body were suitable for pondweed Potamogeton natans L., which was found only in the beaver-created microhabitats.

It is difficult to make any conclusion about the fish distribution in the water body because fishes can use different areas even during the day (Pavlov and Mochek, 2009). The state and structure of the fish population primarily depend on the heterogeneity of habitats of all water bodies within a floodplain (Goetz et al., 2015; Glińska-Lewczuk et al., 2016). Nevertheless, the existence of diverse conditions within a water body is also necessary for normal reproduction (Nunn et al., 2012) because a change in habitat preferences follows a change in food preferences during ontogenesis (Persson and Crowder, 1998). Research on crucian-carp feeding has shown that their fry live almost exclusively in vegetation thickets, and as they switch to benthic food, they prefer deeper places with minimal vegetation (Penttinen and Holopainen, 1992). Pyrzanowski et al. (2021) obtained similar results about weatherfish: small and juvenile fish seek prey in a water column and within macrophytes, while adults prefer benthic food. My results indicate that juvenile fish avoid thickets of emergent plants, but juvenile crucian carp prefer submerged vegetation, which grows in the shore-adjacent area only near burrows.

In our work, almost no significant patterns of tadpole distribution among water body microhabitats were revealed. Only marsh frog larvae avoided burrows in July, preferring more overgrown areas of an unaffected littoral. A possible reason is predator pressure (primarily insects) because this factor can induce a redistribution of larvae within a water body (Kloskowski et al., 2020). Spadefoot tadpoles proved to be dispersed throughout the water body here, but some preferences for BSs were noticed in July. It must be pointed out that between June and July, there was homogenization of conditions in all microhabitats, and differences in depth and plant cover declined. Many neighboring oxbows dried up completely, and all amphibian larvae there died before reaching metamorphosis. This situation became quite common in floodplains in recent years (Antonyuk and Panchenko, 2017; Yermokhin et al., 2018). If tadpoles tend to aggregate near beaver burrows, this phenomenon may be important during shallowing of a water body because such habitats have greater depth and their overgrowth begins later.

Substantial climatic changes have accelerated the succession of ecosystems of floodplain lakes, thus leading to overgrowth, homogenization, and shallowing of water bodies and shortening of the hydroperiod. These processes can cause rapid transformation of water bodies into marshes or meadows. This paper shows that beavers' digging can slow these processes. Even in the absence of these animals, the effects of their past activity affect the present condition of the water body. On the other hand, known conservation strategies for restoration of floodplain water bodies and oxbows usually include expansive and technical approaches: e.g., excavation of channels and lake beds (Buijse et al., 2002; Obolewski et al., 2013). Our results suggest that beaver digging is a natural solution for small water bodies and within protected areas.

Acknowledgments

The author is grateful to Vitaly Osipov for help with the fieldwork and for advice, to Tatiana Gorbushina for assistance with the vegetation surveys, and to Tamara G. Stoyko for help with the identification of the invertebrates. The experiments comply with the current laws of the country in which they were performed.

Appendix A: The list of aquatic plants from the studied water body (Russia, Penza Oblast).

  Control Beaver microhabitats
  June July June July
Spirogyra sp. + – + –
Sparganium erectum L. + – – +
Typha angustifolia L. – + – +
Potamogeton lucens L. – – – +
Potamogeton natans L. – – – –
Potamogeton trichoides Cham. & Schltdl. – + – –
Alisma plantago-aquatica L. + + + +
Alopecurus aequalis Sobol. + + – –
Iris pseudacorus L. – – – –
Ceratophyllum demersum L. – – – –
Callitriche stagnalis Scop. – – – –

Appendix B: The list of taxa of aquatic organisms in the investigated water body (Russia, Penza Oblast). Total numbers of trapped individuals are listed.

  Control Beaver microhabitats
June July June July
Mollusca
Great pond snail Lymnaea stagnalis (Linnaeus, 1758) 86 379 96 239
Marsh pond snail Lymnaea palustris (O.F.Müller, 1774) 6 0 14 1
Great ramshorn Planorbarius corneus (Linnaeus, 1758) 20 14 14 9
Margined ramshorn Planorbis planorbis (Linnaeus, 1758) 8 3 4 0
Crustacea
Tadpole shrimp Lepidurus apus (Linnaeus, 1758) 0 0 1 0
Clam shrimp Cyzicus tetracerus (Krynicki, 1830) 3 0 0 0
Arachnida
Diving bell spider Argyroneta aquatica (Clerck, 1757) 0 0 1 0
Insecta
Emerald dragonfly Cordulia sp. larvae 0 0 1 0
Hawker dragonfly Aeschna sp. larvae 1 0 0 0
Demoiselle damselfly Calopteryx sp. larvae 3 0 1 0
Darter dragonfly Sympetrum sp. larvae 0 0 0 2
Backswimmer bug Notonecta sp. 0 1 0 4
Water bug Naucoris sp. 0 1 6 1
Water stick insect Ranatra sp. 0 0 0 1
Diving beetle Dytiscus sp. larvae 3 0 1 0
Diving beetle Dytiscus sp. imago 2 2 5 4
Water scavenger beetle Hydrophilus sp. larvae 0 0 1 0
Water scavenger beetle Hydrophilus sp. imago 0 0 3 0
Great silver water beetle Hydrous sp. imago 14 4 6 5
Amphibia
Smooth newt Lissotriton vulgaris (Linnaeus, 1758) adults 0 0 1 0
Fire-bellied toad Bombina bombina (Linnaeus, 1761) adults 0 0 2 0
Fire-bellied toad Bombina bombina (Linnaeus, 1761) larvae 3 0 1 0
Spadefoot toad Pelobates vespertinus (Pallas, 1771) larvae 28 129 21 147
Moor frog Rana arvalis Linnaeus, 1758 adults 0 1 0 0
Marsh frog Pelophylax ridibundus (Pallas, 1771) adult 2 8 12 5
Marsh frog Pelophylax ridibundus (Pallas, 1771) larvae 2 15 6 5
Pisces
Weatherfish Misgurnus fossilis (Linnaeus, 1758) adults 0 0 3 0
Weatherfish Misgurnus fossilis (Linnaeus, 1758) juveniles 0 2 0 0
Crucian carp Carassius carassius (Linnaeus, 1758) adults 20 1 0 13
Crucian carp Carassius carassius (Linnaeus, 1758) juveniles 0 21 0 21

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Cite this article as: Bashinskiy IW. 2022. Beaver-created microhabitats in a small water body and their impact on flora and fauna (the Khoper River floodplain, Russia). Int. J. Lim. 58: 16

All Tables

Table 1

The main morphometric characteristics of the studied water body in June–July.

In the text

All Figures

thumbnail Fig. 1

The study area and the location (white circle) of the examined water body (Russia, Penza Oblast). The white line denotes the borders of Privolzhskaya Lesostep Nature Reserve.
In the text
thumbnail Fig. 2

The bathymetric map of the water body in question (June, 2021). The dashed line represents the pre-beaver shoreline (initial perimeter). The hatched areas show beaver-created microhabitats (BS1–BS4). CS1–CS5 are control sites. The red lines indicate littoral zones at distances of 1, 2, 3, and 4 m from the shore.
In the text
thumbnail Fig. 3

Main characteristics of microhabitats of different types (white boxes: control, gray boxes: created by beavers). Depth (a), water temperature (b), and plant cover (c).
In the text
thumbnail Fig. 4

Changes in the number of all plant species (a) and of emergent (b) and submerged plants (c) in microhabitats. White boxes are CSs; gray boxes are BSs.
In the text
thumbnail Fig. 5

A species-conditional triplot based on the canonical correspondence analysis for invertebrate (a: June, c: July) and vertebrate (b: June, d: July) biota. A: adults and imago individuals, J and L: larvae and juvenile individuals. Black dots show BSs, and gray dots represent CSs. SIZE: the size of microhabitat altered by beavers, TEMP: water temperature, COVER: plant cover.
In the text
thumbnail Fig. 6

Relative abundance (a), biomass (b), and the stage of development (Gosner, 1960) (c) of spadefoot tadpoles.
In the text

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