Passive acoustic monitoring as a potential tool to survey animal and ecosystem processes in freshwater environments

1. Biodiversity in freshwater habitats is decreasing faster than in any other type of environment, mostly as a result of human activities. Monitoring these losses can help guide mitigation efforts. In most studies, sampling strategies predominantly rely on collecting animal and vegetal specimens. Although these techniques produce valuable data, they are invasive, time-consuming and typically permit only limited spatial and temporal replication. There is need for the development of complementary methods.
2. As observed in other ecosystems, freshwater environments host animals that emit sounds, either to communicate or as a by-product of their activity. The main freshwater soniferous groups are amphibians, fish, and macroinvertebrates (mainly Coleoptera and Hemiptera, but also some Decapoda, Odonata, and Trichoptera). Biophysical processes such as flow or sediment transport also produce sounds, as well as human activities within aquatic ecosystems.
3. Such animals and processes can be recorded, remotely and autonomously, and provide information on local diversity and ecosystem health. Passive acoustic monitoring (PAM) is an emerging method already deployed in terrestrial environments that uses sounds to survey environments. Key advantages of PAM are its non-invasive nature, as well as its ability to record autonomously and over long timescales. All these research topics are the main aims of ecoacoustics, a new scientific discipline investigating the ecological role of sounds.
4. In this paper, we review the sources of sounds present in freshwater environments. We then underline areas of research in which PAM may be helpful emphasising the role of PAM for the development of ecoacoustics. Finally, we present methods used to record and analyse sounds in those environments.
5. Passive acoustics represents a potentially revolutionary development in freshwater ecology, enabling continuous monitoring of dynamic bio-physical processes to inform conservation practitioners and managers.

     

Acoustic detection and acoustic habitat characterisation of the critically endangered white-bellied heron (Ardea insignis) in Bhutan

1. Growing developmental activities, such as hydropower construction, farm roads, and other human activities, are affecting the critically endangered white-bellied heron (WBH). Out of a known global population of 60, 28 individuals inhabit the river basin area and freshwater lakes and ponds of Bhutan. Several constraints impede continuous monitoring of endangered species, such as the isolated and cryptic nature of the species and the remoteness of its habitat; to date, there are no long-term reference data or techniques implemented for continuous monitoring of this species.
2. In this study, we designed acoustic detection and habitat characterisation methods using long-duration recordings from three habitat areas in Bhutan. Acoustic indices were extracted and used to implement a species-specific call detector and to generate habitat soundscape representations. Using WBH calls annotated in month-long recordings from a known site, a novel indices-based detector was implemented and tested. A total of 960 hr of continuous audio recordings from three habitats in Bhutan were analysed.
3. We found that a species call detector implemented using a combination of acoustic indices (that includes measures of spectral and temporal entropy and different angles of spectral ridges) has a correct detection rate of 81%. Additionally, visual inspection of the species’ acoustic habitat using long-duration false-colour spectrograms enabled qualitative assessment of acoustic habitat structure and other dominant acoustic events.
4. This study proposes a combined approach of species acoustic detection and habitat soundscape analysis for holistic acoustic monitoring of endangered species. As a direct outcome of this work, we documented acoustic reference data on the critically endangered WBH from multiple habitat areas and have analysed its temporal vocalisation patterns across sites.

     

Acoustic monitoring reveals diversity and surprising dynamics in tropical freshwater soundscapes

1. Freshwater systems are globally threatened and in need of enhanced monitoring and assessment. We applied soundscape recording and analysis—which presents an opportunity for long-term, high-resolution animal community monitoring and assessment—to a freshwater context to better understand the acoustic diversity and dynamics of these systems.
2. We recorded the aquatic soundscape of a Neotropical freshwater swamp in Costa Rica for 23 days in January and February 2015 during the dry season. We classified biological sound types in these recordings and developed measurements of richness and occupancy based on this classification. We also calculated six complementary acoustic indices to assess soundscape diversity and daily and longer-term soundscape dynamics, and we examined correlations between these acoustic indices and sound type metrics.
3. We found rich soundscapes in which biological sounds were almost always present, and we classified 18 sound types that we attribute to aquatic insects. These sound types showed distinct daily patterns and exhibited temporal and spectral acoustic niche partitioning. Sound type richness was most correlated with the number of peaks index (correlation = .36; p < .001), while sound type occupancy was most correlated with the Bioacoustic Index (correlation = .92; p < .001). In contrast to generally high levels of acoustic activity, there were brief (approximately 1 hr), unexpected quiet periods around dawn and dusk.
4. This study represents an early attempt to comprehensively describe tropical freshwater soundscapes in a systematic and quantitative manner. We demonstrate that sound type classification and the quantification of acoustic occupancy capture aspects of soundscape diversity and dynamics that are complementary to those assessed by acoustic indices. Our analyses reveal that the soundscapes of this tropical wetland were diverse and exhibited daily dynamics that differed from those found in other ecosystems.

     

Monitoring the acoustic activity of an aquatic insect population in relation to temperature,vegetation and noise

1. Acoustic population monitoring is a noninvasive method that can be deployed continuously over long periods of time and at large spatial scales. One of the newly discovered threats acting on biological diversity is anthropogenic noise. High levels of anthropogenic noise occur in aquatic environments, yet their effects on animals living in freshwater habitats have very rarely been investigated.
2. Here, we used acoustic monitoring and automatic detection to assess the acoustic activity of a population of a soniferous freshwater insect.
3. The sounds emitted by the corixid Micronecta scholtzi were recorded in a Mediterranean pond with an array of 12 hydrophones. An automatic analysis based on a measure of the amplitude found in the frequency band of M. scholtzi was developed to assess the level of acoustic activity. We used functional linear models, accounting for the periodicity of the calling behaviour, to estimate the possible effect of temperature, vegetation and a noise due to an immersed engine.
4. The automatic analysis was validated as an efficient method to measure the acoustic activity. The monitoring revealed a clear 24-hr pattern in the acoustic activity of M. scholtzi and three peaks of activity during the morning. Functional linear models revealed negative effects of both temperature and vegetation and showed that an engine noise, played back for 2 hr during the night, elicited an increase in the level of acoustic activity of the population. Moreover, a cross-correlation procedure showed that noise delayed the acoustic activity of the population.
5. Our results suggest that acoustic survey and automatic detection are efficient methods to monitor the acoustic activity of an insect population especially in response to an anthropogenic disturbance.

     

Six steps towards operationalising freshwater ecoacoustic monitoring

1. Applications in bioacoustics and its sister discipline ecoacoustics have increased exponentially over the last decade. However, despite knowledge about aquatic bioacoustics dating back to the times of Aristotle and a vast amount of background literature to draw upon, freshwater applications of ecoacoustics have been lagging to date.
2. In this special issue, we present nine studies that deal with underwater acoustics, plus three acoustic studies on water-dependent birds and frogs. Topics include automatic detection of freshwater organisms by their calls, quantifying habitat change by analysing entire soundscapes, and detecting change in behaviour when organisms are exposed to noise.
3. We identify six major challenges and review progress through this special issue. Challenges include characterisation of sounds, accessibility of archived sounds as well as improving automated analysis methods. Study design considerations include characterisation analysis challenges of spatial and temporal variation. The final key challenge is the so far largely understudied link between ecological condition and underwater sound.
4. We hope that this special issue will raise awareness about underwater soundscapes as a survey tool. With a diverse array of field and analysis tools, this issue can act as a manual for future monitoring applications that will hopefully foster further advances in the field.

     

Singing streams: Describing freshwater soundscapes with the help of acoustic indices

1. Understanding soundscapes, that is, the totality of sounds within a location, helps to assess nature in a more holistic way, providing a novel approach to investigating ecosystems. To date, very few studies have investigated freshwater soundscapes in their entirety and none across a broad spatial scale.
2. In this study, we recorded 12 freshwater streams in South East Queensland continuously for three days and calculated three acoustic indices for each minute in each stream. We then used principal component analysis of summary statistics for all three acoustic indices to investigate acoustic properties of each stream and spatial variation in their soundscapes.
3. All streams had a unique soundscape with most exhibiting diurnal variation in acoustic patterns. Across these sites, we identified five distinct groups with similar acoustic characteristics. We found that we could use summary statistics of AIs to describe daytimes across streams as well. Most difference in stream soundscapes was observed during the daytime with significant variation in soundscapes both between hours and among sites.
4. Synthesis and Application. We demonstrate how to characterize stream soundscapes by using simple summary statistics of complex acoustic indices. This technique allows simple and rapid investigation of streams with similar acoustic properties and the capacity to characterize them in a holistic and universal way. While we developed this technique for freshwater streams, it is also applicable to terrestrial and marine soundscapes.

     

Ecoacoustics can detect ecosystem responses to environmental water allocations

1. Globally, water abstraction for human consumption and irrigated agriculture leads to significant changes in aquatic ecosystems. To counter these detrimental effects, water releases—often termed environmental water allocations—restore overbank flow or are delivered to artificially disconnected wetlands. While a suite of monitoring methods is available, few programmes track continuous change in biota, mainly because repeated remote site visits can be prohibitively expensive.
2. In this paper, we propose a new approach to environmental flow monitoring, using ecoacoustic methods. We test acoustic monitoring of frog and waterbird responses to environmental water deliveries in the Goulburn Broken, a valley in the southern Murray–Darling river system. Response to three major environmental water deliveries within 2 years was monitored at four sites along Reedy swamp. Every 2 weeks, 30 s were recorded every 30 min, for a total of 24 hr.
3. We used two analysis strategies—manual counts of bird calls, as well as ecoacoustic indices, which describe the sonic properties of the acoustic spectrum at a site. Manual counts demonstrated that water-dependent birds were clearly responding to environmental water deliveries, whereas non-water-dependent species did not show any increases in activity. After restricting the analysis to the dawn chorus of birds and frogs, two acoustic indices (the median amplitude and the acoustic complexity index) showed responses to watering events.
4. Ecoacoustic methods show promise for continuous response monitoring to environmental water allocations. However, the first strategy—manual annotation of calls—might be too labour intensive for standard monitoring programmes. The second strategy—index-based approaches—can also detect ecological responses, although further investigation using control sites is needed. Automated call classifiers are an alternative that is currently being developed for endangered species. We also encourage simultaneous monitoring of the soundscape above and under water.

     

Assessing the value of acoustic indices to distinguish species and quantify activity: A case study using frogs

1. Soundscapes can provide information about a wide range of habitats and species through the recording of vocalisations over long temporal scales. Because of the large volumes of data collected, computational approaches, such as the application of acoustic indices, are required to extract useful information from long-duration recordings.
2. Acoustic indices summarise various soundscape features into frequency ranges over defined time intervals and can aid in the visual exploration, detection, and analysis of species vocalisation patterns. Here, we examine the performance of combinations of three acoustic indices commonly used in visual exploration, the acoustic complexity index, the temporal entropy spectrum index, and the event spectrum index, and assess their ability to distinguish species and describe acoustic features commonly used to detect species and analyse activity. Our case study focuses on three frog species with distinct call structures from Bickerton Island, Northern Territory, Australia. Call structure was categorised based on the number of pulses and harmonics.
3. We summarised acoustic activity by calculating acoustic indices in 256 equal-sized bins over the entire the frequency spectrum, for 30-s intervals, and found that acoustic index values could be used to distinguish species and describe acoustic features. The acoustic complexity index was the most effective index for distinguishing species. To describe acoustic features, we examined correlations between acoustic index values and summarised acoustic features, including call rate, total duration, loudness and signal-to-noise ratio. In single-pulsed species with no harmonics, we found spectral index values were significantly and sometimes strongly correlated with acoustic features. In comparison, species with harmonics were found to be weakly and less frequently correlated with acoustic features even if sampled calls were loud and have high signal-to-noise ratio. We suggest that acoustic indices have the potential to describe acoustic features in single-pulsed species but are limited in those with harmonics.
4. We conclude that acoustic indices can be a useful tool to distinguish some anuran species and to broadly understand specific acoustic features used to analyse calling activity over long periods of time.
5. Further research is required to better understand the relationships between acoustic indices and acoustic features to determine the general utility of indices to detect and distinguish audible species and to identify other acoustic features of various taxa.

     

Temporally adaptive acoustic sampling to maximize detection across a suite of focal wildlife species

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Looking for love under the ice: Using passive acoustics to detect burbot (Lota lota: Gadidae) spawning activity

1. Burbot (Lota lota: Gadidae) is a difficult species to manage effectively due to its preference for deep-water habitats and under-ice spawning behaviour, resulting in a poor understanding of its reproductive activity. However, the use of acoustic signalling by burbot as part of their mating system has recently been described and this behaviour may provide a means of investigating questions regarding the spatial and temporal distribution of spawning aggregations using passive acoustic monitoring.
2. We used audio and video recording to confirm that burbot vocalise and that these vocalisations can be detected under field conditions as well as to characterise the relationship between burbot acoustic signalling and spawning behaviour. We also evaluated the feasibility of locating and monitoring burbot spawning aggregations in real time using passive acoustics.
3. Burbot vocalisations were difficult to identify with only about 6% of the recordings containing calls being successfully identified as such in the field. Burbot vocalised more often between sundown and sunrise than during daylight hours. Calls recorded at night tended to be lower frequency, longer duration, and have lower bandwidth than those made during the day.
4. Burbot vocalisations could not be recorded in conjunction with video recordings of spawning activity, indicating that burbot may not call during active spawning, but may use acoustic communication to signal the onset of reproductive readiness and to form pre-spawning aggregations.
5. While burbot calls were readily identifiable, observers had a difficult time identifying burbot calls in real time under field conditions. Passive acoustic monitoring demonstrates considerable potential as a management tool to locate burbot spawning grounds and identify periods of activity, but may not be an appropriate technique for monitoring spawning activity in real time.

     

Diurnal variation in freshwater ecoacoustics: Implications for site-level sampling design

1. Ecoacoustic methods are increasingly used to monitor the state of populations and ecosystems. In freshwater environments, they present the clear advantages of being non-invasive, reducing bias, and providing continuous observations instead of only limited sampling snapshots in time. However, similar to standard bioassessment methods, temporal variation and choice of indicators can greatly influence ecoacoustic assessments, highlighting the importance of sampling and analysis design.
2. In this study, we quantified diurnal variation in underwater sound and its effect on sampling regimes for two waterholes in the Einasleigh River, Northern Australia. Recording continuously for 6 days, and subsampling 5 s every 10 min, we found 22 distinct sounds that were emitted by fish, Hemiptera and Coleoptera as well as another 22 of abiotic or unknown origin.
3. Through rarefaction analyses, we found that subsampling the data to 60% of the recorded sound events resulted in capture of most of the 44 identified sound types. Temporal heterogeneity—patchy sound events through time—needs to be considered when maximising detected sound events. Reducing the sampling interval from every 10 min to half-hourly or hourly had a much greater effect on capturing all sound types compared to the number of days recorded or the length of the recording. Overall, only 10–20% of the sound events need to be annotated for most sound types to be described; for example, restricting analysis of the days recorded to only three and the recording interval to 0.5–1 s. Acoustic indices were dominated by three main event types—a diurnally flowing creek, a nocturnal chorus of Hemiptera, as well as a dawn chorus of terapontid fishes.
4. We conclude with two key messages: First, a select group of informative signals can be monitored using very simple methods—namely, converting an audio stream into indices using freely available software. Second, however, to detect less acoustically dominant sound events, manual annotation or single call processing will still be needed. While these findings are encouraging, similar analysis will need to be conducted within other freshwater ecosystems before general conclusions about optimal sampling regimes can be drawn.

     

SILIC: A cross database framework for automatically extracting robust biodiversity information from soundscape recordings based on object detection and a tiny training dataset

• 1.Passive acoustic monitoring (PAM) offers many advantages comparing with other survey methods and gains an increasing use in terrestrial ecology, but the massive effort needed to extract species information from a large number of recordings limits its application. The convolutional neural network (CNN) has been demonstrated with its high performance and effectiveness in identifying sound sources automatically. However, requiring a large amount of training data still constitutes a challenge.
• 2.Object detection is used to detect multiple objects in photos or videos and is effective at detecting small objects in a complex context, such as animal sounds in a spectrogram and shows the opportunity to build a good performance model with a small training dataset. Therefore, we developed the Sound Identification and Labeling Intelligence for Creatures (SILIC), which integrates online animal sound databases, PAM databases and an object detection-based model, for extracting information on the sounds of multiple species from complex soundscape recordings.
• 3.We used the sounds of six owl species in Taiwan to demonstrate the effectiveness, efficiency and application potential of the SILIC framework. Using only 786 sound labels in 133 recordings, our model successfully identified the species' sounds from the recordings collected at five PAM stations, with a macro-average AUC of 0.89 and a mAP of 0.83. The model also provided the time and frequency information, such as the duration and bandwidth, of the sounds.
• 4.To our best knowledge, this is the first time that the object detection algorithm has been used to identify sounds of multiple wildlife species. With an online sound-labeling platform embedded and a novel data preprocessing approach (i.e., rainbow mapping) applied, the SILIC shows its good performance and high efficiency in identifying wildlife sounds and extracting robust species, time and frequency information from a massive amount of soundscape recordings based on a tiny training dataset acquired from existing animal sound databases. The SILIC can help expand the application of PAM as a tool to evaluate the state of and detect the change in biodiversity by, for example, providing high temporal resolution and continuous information on species presence across a monitoring network.

     

Pond Acoustic Sampling Scheme: A draft protocol for rapid acoustic data collection in small waterbodies

1. Freshwater conservation is vital to the maintenance of global biodiversity. Ponds are a critical, yet often under‐recognized, part of this, contributing to overall ecosystem functioning and diversity. They provide habitats for a range of aquatic, terrestrial, and amphibious life, often including rare and declining species.
2. Effective, rapid, and accessible survey methods are needed to enable evidence‐based conservation action, but freshwater taxa are often viewed as “difficult”—and few specialist surveyors are available. Datasets on ponds are therefore limited in their spatiotemporal coverage.
3. With the advent of new recording technologies, acoustic survey methods are becoming increasingly available to researchers, citizen scientists, and conservation practitioners. They can be an effective and noninvasive approach for gathering data on target species, assemblages, and environmental variables. However, freshwater applications are lagging behind those in terrestrial and marine spheres, and as an emergent method, research studies have employed a multitude of different sampling protocols.
4. We propose the Pond Acoustic Sampling Scheme (PASS), a simple protocol to allow a standardized minimal sample to be collected rapidly from small waterbodies, alongside environmental and methodological metadata. This sampling scheme can be incorporated into a variety of survey designs and is intended to allow access to a wide range of participants, without requiring complicated or prohibitively expensive equipment.
5. Adoption of this sampling protocol would enable consistent sound recordings to be gathered by researchers and conservation organizations, and allow the development of landscape‐scale surveys, data sharing, and collaboration within an expanding freshwater ecoacoustic community—rather than individual approaches that produce incompatible datasets. The compilation of standardized data would improve the prospects for effective research into the soundscapes of small waterbodies and aid freshwater conservation efforts.

     

Environmental heterogeneity influences life-form richness and species composition but not species richness of aquatic macrophytes in tropical coastal rivers

1. The coastal rivers of the São Paulo State in SE Brazil have different lengths and seawater influence. We evaluated whether: (1) environmental heterogeneity (EH) is associated with the species and life-form richness of aquatic macrophytes; and (2) EH and geographical distance influence species composition in these coastal rivers.
2. We recorded the macrophyte species and life form occurrence and collected explanatory variables characterising the water, sediment, and river channel at 100 sampling sites over 8 rivers. We applied a principal component analysis to the explanatory variables and calculated the rivers' EH using the range of principal component 1 scores. We also determined the position of each river mouth along the coastline to measure the distance between the rivers. We used quasi-Poisson generalised linear models to evaluate the effects of EH on richness of species and life forms. To determine the effect of EH and geographical distance (Euclidean distance matrices) on the variation in species composition (Jaccard dissimilarity matrix) among the rivers, we applied multiple regressions on distance matrices.
3. The most heterogeneous river had heterogeneity score about five times greater than the least heterogeneous river. Sediment salinity, river width, total phosphorus concentration of water and distance from river mouth were the most important variables contributing to the rivers' EH. We found that EH did not explain variation in species richness; however, it had a significant positive relationship with life-form richness. The effect of EH was greater than that of the geographical distance on the variation in species composition among the rivers. The pairs of rivers with the most similar EH were the most similar in species composition, but not all of them were geographically close.
4. We conclude that EH influences life-form richness but does not influence species richness of aquatic macrophytes in the coastal rivers we studied; however, EH does influence species composition regardless of geographical distance among rivers.

     

Interspecific variation in the response of fish to anthropogenic noise

1. Elevated levels of anthropogenic noise, especially those observed through boating activity, can negatively impact fish species, but it remains unclear which species are most affected and which behavioural metrics are best used in assessing fish responses to underwater noise. The effects of boat sounds on freshwater species are of particular interest because freshwater environments are less studied than the marine realm despite comparably high levels of biodiversity.
2. In the current study, we examine the behavioural responses to boat noise in two freshwater species that differ in their hypothesised response to sound inputs: the spottail shiner (Notropis hudsonius), a species with known hearing specialisations, and the bluegill sunfish (Lepomis macrochirus), a species with more generalised hearing capabilities. Fish were presented with boat noise in a laboratory setting, and their swimming, escape and foraging behaviours were assessed to examine differential responses in relation to hypothesised hearing abilities.
3. Both species showed a decrease in general swimming behaviours but an increase in erratic movements in response to boat noise, indicative of stress responses for both species. Despite the similarities in response based on swimming behaviours however, only spottail shiners exhibited true escape responses to the onset of the noise stimulus, suggesting a more extreme reaction in the species with a more refined hearing ability.
4. Taken together, these results show that freshwater fish can respond to increased levels of anthropogenic noise, but that the severity of the response may differ based on auditory structures and therefore presumed hearing ability. The differences seen between behavioural metrics used (swimming vs. escape responses) also demonstrate how care must be taken in choosing a metric when developing exposure guidelines for underwater sound exposures, as different metrics could lead to differential impact assessments.

     

Phylogenetic placement of environmental sequences using taxonomically reliable databases helps to rigorously assess dinophyte biodiversity in Bavarian lakes (Germany)

1. Reliable determination of organisms is a prerequisite to explore their spatial and temporal occurrence and to study their evolution, ecology, and dispersal. In Europe, Bavaria (Germany) provides an excellent study system for research on the origin and diversification of freshwater organisms including dinophytes, due to the presence of extensive lake districts and ice age river valleys. Bavarian freshwater environments are ecologically diverse and range from deep nutrient-poor mountain lakes to shallow nutrient-rich lakes and ponds.
2. We obtained amplicon sequence data (V4 region of small subunit-rRNA, c. 410 bp long) from environmental samples collected at 11 sites in Upper Bavaria. We found 186 operational taxonomic units (OTUs) associated with Dinophyceae that were further classified by means of a phylogenetic placement approach.
3. The maximum likelihood tree inferred from a well-curated reference alignment comprised a systematically representative set of 251 dinophytes, covering the currently known molecular diversity and OTUs linked to type material if possible. Environmental OTUs were scattered across the reference tree, but accumulated mostly in freshwater lineages, with 79% of OTUs placed in either Apocalathium, Ceratium, or Peridinium, the most frequently encountered taxa in Bavaria based on morphology.
4. Twenty-one Bavarian OTUs showed identical sequences to already known and vouchered accessions, two of which are linked to type material, namely Palatinus apiculatus and Theleodinium calcisporum. Particularly within Peridiniaceae, delimitation of Peridinium species was based on the intraspecific sequence variation.
5. Our approach indicates that high-throughput sequencing of environmental samples is effective for reliable determination of dinophyte species in Bavarian lakes. We further discuss the importance of well-curated reference databases that remain to be developed in the future.

     

Lessons from the macroinvertebrates: species‐genetic diversity correlations highlight important dissimilar relationships

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Salinisation reduces biodiversity in neighbouring freshwater patches in experimental metacommunities

1. Aquatic ecosystems are biodiversity hot spots across many landscapes; therefore, the degradation of these habitats can lead to decreases in biodiversity across multiple scales. Salinisation is a global issue that threatens freshwater ecosystems by reducing water quality and local biodiversity. The effects of salinity on local processes have been studied extensively; however, the effects of salinisation or similar environmental stressors within a metacommunity (a dispersal network of several distinct communities) have not been explored.
2. We tested how the spatial heterogeneity and the environmental contrast between freshwater and saline habitat patches influenced cladoceran biodiversity and species composition at local and regional scales in a metacommunity mesocosm experiment. We defined spatial heterogeneity as the proportion of freshwater to saltwater patches within the metacommunity, ranging from a freshwater-dominated metacommunity to a saltwater-dominated metacommunity. Environmental contrast was defined as the environmental distance between habitat patches along the salinity gradient in which low-contrast metacommunities consisted of freshwater and low-salinity patches and high-contrast metacommunities consisted of freshwater and high-salinity patches.
3. We hypothesised that the α-richness of freshwater patches and metacommunity γ-richness would decrease as freshwater patches became less abundant along the spatial heterogeneity gradient in both low- and high-contrast metacommunities, because there would be fewer freshwater patches that could serve as source populations for declining populations. We hypothesised that low-contrast metacommunities would support more species across the spatial heterogeneity gradient than high-contrast metacommunities, because, via dispersal, low-salinity patches can support halotolerant freshwater species that can mitigate population declines in neighbouring freshwater patches, whereas` high-salinity patches will mostly support halophilic species, providing fewer potential colonisers to freshwater patches.
4. We found that α-richness of freshwater mesocosms and metacommunity γ-richness declined in saline-dominated metacommunities regardless of the environmental contrast between the freshwater and saline mesocosms. We found that environmental contrast influenced freshwater and saline community composition in low-contrast metacommunities by increasing the abundances of species that could tolerate low-salinity environments through dispersal, whereas freshwater and high-salinity communities showed limited interactions through dispersal.
5. Freshwater mesocosms had a disproportionate effect on the local and regional biodiversity in these experimental metacommunities, indicating that habitat identity may be more important than habitat diversity for maintaining biodiversity in some metacommunities. This study further emphasises the importance in maintaining multiple species-rich habitat patches across landscapes, particularly those experiencing landscape-wide habitat degradation.