Wetland Invertebrates Research Articles

Pollution-tolerant invertebrates enhance greenhouse gas flux in urban wetlands.

One of the goals of urban ecology is to link community structure to ecosystem function in urban habitats. Pollution-tolerant wetland invertebrates have been shown to enhance greenhouse gas (GHG) flux in controlled laboratory experiments, suggesting that they may influence urban wetland roles as sources or sinks of GHG. However, it is unclear if their effects can be detected in highly variable conditions in a field setting. Here we use an extensive data set on carbon dioxide (CO2 ), methane (CH4 ), and nitrous oxide (N2 O) flux in sediment cores (n=103) collected from 10 urban wetlands in Melbourne, Australia during summer and winter in order to test for invertebrate enhancement of GHG flux. We detected significant multiplicative enhancement effects of temperature, sediment carbon content, and invertebrate density on CH4 and CO2 flux. Each doubling in density of oligochaete worms or large benthic invertebrates (oligochaete worms and midge larvae) corresponded to ~42% and ~15% increases in average CH4 and CO2 flux, respectively. However, despite exceptionally high densities, invertebrates did not appear to enhance N2 O flux. This was likely due to fairly high organic carbon content in sediments (range 2.1-12.6%), and relatively low nitrate availability (median 1.96μmol/L NO3- -N), which highlights the context-dependent nature of community structural effects on ecosystem function. The invertebrates enhancing GHG flux in this study are ubiquitous, and frequently dominate faunal communities in impaired aquatic ecosystems. Therefore, invertebrate effects on CO2 and CH4 flux may be common in wetlands impacted by urbanization, and urban wetlands may make greater contributions to the total GHG budgets of cities if the negative impacts of urbanization on wetlands are left unchecked.

Transferability of bioassessment indices among water body types and ecoregions: A California experiment in wetland assessment

Biological assessment of aquatic resources requires the availability of bioassessment tools that work in all waterbody types and regions of interest. Developing new assessment tools may require several years of data collection and substantial investment of resources, which may not be an option for some aquatic resource managers. Adapting tools developed for different regions or wetland types may be an attractive alternative to developing new indices, provided they work well in the novel setting. In this study, we explore the transferability of two bioassessment indices for application to depressional wetlands in California, which are wetland type of management concern but for which bioassessment tools don’t currently exist. We tested the applicability of a depressional wetland invertebrate index of biotic integrity (IBI) developed in the San Francisco Bay region of northern California for application in the drier regions of southern California (i.e. geographic transferability), and the ability to apply a riverine benthic diatom IBI to benthic diatoms in depressional wetlands (i.e. water body type transferability). We evaluated the accuracy and responsiveness of the existing Indices for use in depressional wetlands and refined reference definitions and recalibrated thresholds relative to stressor gradients to maximize index performance. Performance of the adapted indices was compared to that of an existing habitat assessment tool (the California Rapid Assessment Method; CRAM) that has been developed for statewide application of depressional wetlands. Finally, we demonstrate application of the revised indices for ambient assessment of depressional wetland condition in southern California. Recalibrating both the macroinvertebrate and diatom indices to reference thresholds based on nutrient concentrations resulted in lower coefficient of variation among reference sites, greater differentiation between reference and non-reference and stronger relationship with stressors than when reference thresholds were based on landscape disturbance. Overall, the simple adjustment of the reference definition allowed us to transfer the indices with no structural changes to the metrics. This approach can facilitate future index adaptations that allow practitioners to include waterbody types for which there is no current index into routine biomonitoring programs.

Effects of spatial scale and habitat on the diversity of diapausing wetland invertebrates

Effects of spatial scale and habitat on the diversity of diapausing wetland invertebrates

Connecting the trophic dots: responses of an aquatic bird species to variable abundance of macroinvertebrates in northern boreal wetlands

To evaluate variation in abundance of boreal wetland macroinvertebrates and test for effects of this variation on the diet and habitat use of a bird species that consumes aquatic invertebrates (lesser scaup, Aythya affinis), we collected macroinvertebrates and birds and conducted bird surveys (June–August) at two wetland complexes in northwestern Canada and assessed diet composition using an isotopic approach. At both study areas, for macroinvertebrate taxa reported to be key prey items for scaup, biomass varied intra-seasonally and annually, but patterns differed among taxa and between areas. Macroinvertebrate biomass varied strongly across wetlands within study areas, and isotopic mixing models indicated that local heterogeneity in macroinvertebrate biomass was reflected in duckling diets, which also varied across wetlands, indicating a generalist foraging strategy for this species. Wetland habitats used by brood-rearing female scaup had greater amphipod and gastropod biomasses. Our results show that boreal wetland macroinvertebrate abundances vary considerably across coarse and fine spatial scales. Female scaup and their ducklings appear well adapted to exploit this dynamic food resource, but overall productivity of scaup may depend on the abundances of certain taxa, suggesting that conservation efforts should focus on maintaining abundant populations of key wetland invertebrates.

Impacts of terrestrial habitat transformation on temporary wetland invertebrates in a sclerophyllous Sand fynbos landscape

There has been recent debate about the extent to which human disturbance of the landscape affects wetland invertebrates given that the organisms are already adapted to high levels of natural disturbance. Using repeated sampling of a set of 12 temporary wetlands occurring in a differentially transformed Sand fynbos landscape in Cape Town (South Africa), we investigated patterns of macroinvertebrate and microcrustacean assemblage composition, richness and diversity in relation to a physico-chemical gradient resulting from variable habitat loss in the adjacent landscape. Both macroinvertebrates and microcrustaceans showed clear gradational changes in assemblage composition in relation to the surrounding cover of indigenous vegetation (as a proxy for habitat loss). Although the composition of assemblages appeared to be affected by this gradient of habitat transformation, no relationships were detected using various measures of taxon richness and diversity. At the small spatial scale for which patterns are analysed, the influence of natural variation on invertebrate assemblages appears to have been overridden by that resulting from habitat transformation in the adjacent landscape. Depression wetlands embedded in Sand fynbos habitat appear to be unique in terms of their physico-chemistry (acidic and rich in humic substances) and the invertebrate assemblages they support.

Inorganic and organic contaminants in Alaskan shorebird eggs.

Many shorebird populations throughout North America are thought to be declining, with potential causes attributed to habitat loss and fragmentation, reduced prey availability, increased predation, human disturbance, and increased exposure to environmental pollutants. Shorebirds may be particularly vulnerable to contaminant exposure throughout their life cycle, as they forage primarily on invertebrates in wetlands, where many contaminants accumulate disproportionately in the sediments. Therefore, it is important to document and monitor shorebird populations thought to be at risk and assess the role that environmental contaminants may have on population declines. To investigate potential threats and provide baseline data on shorebird contaminant levels in Alaskan shorebirds, contaminant concentrations were evaluated in shorebird eggs from 16 species residing in seven geographic distinct regions of Alaska. Similar to previous studies, low levels of most inorganic and organic contaminants were found, although concentrations of several inorganic and organic contaminants were higher than those of previous studies. For example, elevated strontium levels were observed in several species, especially black oystercatcher (Haematopus bachmani) sampled in Prince William Sound, Alaska. Additionally, contaminant concentrations varied among species, with significantly higher concentrations of inorganic contaminants found in eggs of pectoral sandpiper (Calidris melanotos), semipalmated sandpiper (Calidris pusilla), black oystercatcher, and bar-tailed godwit (Limosa lapponica). Similarly, significantly higher concentrations of some organic contaminants were found in the eggs of American golden plover (Pluvialis dominica), black-bellied plover (Pluvialis squatarola), pacific golden plover (Pluvialis fulva), bar-tailed godwit, and semipalmated sandpiper. Despite these elevated levels, current concentrations of contaminants in shorebird eggs suggest that breeding environments are relatively free of most contaminants and that contaminant concentrations are below levels (except potentially strontium) that would likely affect the survival of individuals and consequently regulate the species at the population level.

No association between the use of Bti for mosquito control and the dynamics of non-target aquatic invertebrates in French coastal and continental wetlands

The environmental safety of Bacillus thuringiensis subsp. israelensis (Bti) is still controversial, mainly because most of the previous field studies on its undesired effects were spatially limited and did not address the relationship between community similarity and application time and frequency. No general statement can therefore be drawn on the usage conditions of Bti that insure protection of non-target organisms.The present study was conducted in eight sites distributed over the main geographical sectors where mosquito control is implemented in mainland France and Corsica. Changes in non-target aquatic invertebrates were followed at elapsed time after repeated applications of two Bti formulations (VectoBac® WDG or 12AS) up to four consecutive years. We examined the influence of both larvicide treatments and environmental variables on community dynamics and dissimilarity between treated and control areas. As it can be argued that chironomids are the most vulnerable group of non-target invertebrates, we scrutinised potential Bti-related effects on the dynamics of their community.The use of VectoBac® WDG and 12AS in coastal and continental wetlands had no immediate or long-term detectable effect on the taxonomic structure and taxa abundance of non-target aquatic invertebrate communities, including chironomids. This applied to the main habitats where mosquito larvae occur, regardless of their geographic location. Flooding, whose frequency and duration depend on local meteorological and hydrological conditions, was identified as the main environmental driver of invertebrate community dynamics.Our findings add support to the environmental safety of currently available Bti formulations when following recommended application rates and best mosquito control practices.

Fringing trees may provide a refuge from prolonged drying for urban wetland invertebrates

Climate change is causing prolonged drying in many seasonal wetlands, including urban wetlands, potentially affecting aquatic invertebrates that take refuge in wetland sediment during dry periods and thereby threatening wetland biodiversity. We collected sediment from two habitats: open water (OW) and fringing trees (FT), in eight urban wetlands after seasonal inundation had ended. Both habitats are inundated during winter–spring and dry in summer–autumn. Each sediment sample was divided into subsamples. One set of subsamples were inundated in the laboratory to test the hypothesis that emerging invertebrate assemblages would differ between OW and FT sediments. Another set of subsamples was dried, stored for a year, and inundated to test the hypothesis that prolonged drying would reduce the abundance and taxa richness of emerging invertebrates. The composition of emerging invertebrate assemblages differed between habitats, with more amphibious species found in FT sediment. Invertebrate responses to prolonged drying and storage varied among species: for some, effects depended on habitat type, while others delayed emergence or showed no response. Microcrustacean abundance was unaffected by drying, suggesting that their productivity during refilling may resist drier water regimes. Surface temperatures of dry sediment are cooler beneath FT, and this sediment has higher organic matter, holds more water and is less dense than OW sediment; and FT sediment remained cooler than OW sediment in the laboratory, despite the absence of shading. Fringing trees may therefore provide a refuge for some freshwater invertebrates relying on dormant stages in the sediment to survive drying in urban wetlands.

Invertebrate community turnover following control of an invasive weed

Non-native invasive plants are a significant threat to native biodiversity as they change community structure and ecosystem processes and it is becoming imperative that these invasive plants are controlled. We investigated the response of wetland invertebrates to glyphosate chemical control of an introduced tree, grey willow (Salix cinerea), in a before-after-control-impact (BACI) experiment within a New Zealand wetland. Our findings suggest that control of grey willow resulted in significant increases in the abundance of invertebrates within 2 years after herbicide application. Analysis of beetle communities revealed that the abundance and species richness of herbivorous and predacious beetles, the abundance of introduced species, and the species richness of native and introduced species were all significantly higher in the herbicide-treated plots than in the unsprayed plots. The composition of the beetle community separated into two clear groups—plots before spray/no spray and plots 1 and 2 years after herbicide application. We suggest that the beetle compositional changes were driven by complex changes in vegetation associated with the canopy collapse of grey willow trees and the initial influx of introduced plant species, predominantly weedy annuals and perennials, after willow control. Restoration via invasive plant control can promote the reestablishment of invertebrate communities typical of native wetlands, but their long-term sustainability is contingent on prevention of grey willow reinvasion and reestablishment of the native plant habitat.

Spatial distribution of resting stages of crustaceans and other aquatic invertebrates in playa wetlands

Crustaceans in temporary wetlands have resting stages that allow them to persist in the soil during dry phases. We examined the spatial distribution of viable resting stages of invertebrates in the soil of playa wetlands. We took soil cores near the center and near the edge of the playas and in the surrounding upland of the dry playas. We then rehydrated the soil cores for two weeks. We found higher abundance and taxa richness, and a different taxonomic composition in the playa center than in the upland. Invertebrate abundance and taxa richness were intermediate at the wetland edge and taxonomic composition overlapped with that from the center and edge. Crustaceans (tadpole shrimps and calanoid copepods) were more abundant in the wetland center than from the edge or upland. Environmental factors and invertebrate behavior may play a role in organizing the spatial distribution of aestivating stages of aquatic invertebrates.

Needle in a Haystack: Predicting the Occurrence of Wetland Invertebrates on the Basis of Simple Geographical Data. A Case Study on Two Threatened Micro-Mollusc Species (Gastropoda: Vertiginidae) from Poland

The aim of this study was to evaluate whether simple geographical data can be used to predict the probability of the presence of two threatened snail species: Vertigo angustior and V. moulinsiana, occurring within wetland areas across their central European range. Our research was based on data from 226 localities across Poland fitting with the species requirements according to the current literature. We used Canonical Variance Analysis with variance partitioning and tree classifications to model relationships between the species presence/absence and rough geographic features describing each locality (landscape and geomorphological types and hydrography). Our results have shown that the localities of the studied species are related to particular types of landform and landscape and are not distributed uniformly. Both species occur predominantly in young postglacial lowland landscapes and are usually associated with particular succession stages of lakes, with V. moulinsiana occurring in earlier phases of habitat evolution. Results from modelling show that cost- and time effectiveness of surveying for these species could be substantially increased by avoiding particular location types. The prediction models presented will be a useful tool for planning future surveys of the vertiginids and in studies on their ecology and distribution.

The impact of anthropogenic noise on wetland habitats

Impacts of anthropogenic noise is an important but poorly studied aspect of wetland ecology. Many aquatic animals use sound to attract mates, coordinate movements, defend territory, and detect predators. However, anthropogenic noise (e.g., from boat or automobile motors) are increasingly important in wetlands. Therefore, it is important to understand how anthropogenic noise pollution affects these habitats. I sampled wetland soundscapes in urbanized and rural areas in northeastern Ohio, and used Raven software to analyze the data. Above the water line, average sound power was 38.9db–73.9db, which was similar to below the water line (48.6db–78.5db). Frequencies in the band 0–5 kHz had the highest overall power both above and below the water. One of the most urbanized sites had above/below readings as high as 83.9/79.3 dB. In comparison, one of the most rural sites had readings as low as 47.6/66.8 dB. At both urbanized and rural sites, the 0–5 kHz band had the highest overall power. I further tested if wetland invertebrates could be impacted by anthropogenic sounds. I found that Procambarus acutus crayfish produced audible clicks (frequency range 6 kHz–45 kHz, average power 69–73.5 dB). Ongoing laboratory research is testing if motorboat noise affects P. acutus behavior and sound production.

Response of Wetland Invertebrate Communities to Local and Landscape Factors in North Central Oklahoma

Invertebrates play an important role influencing wetland functions. Specifically, they provide important food for many waterbirds and other wetland species. To better understand the role environmental factors play in influencing invertebrates, we examined the influence of local and landscape factors on invertebrate communities inhabiting depressional wetlands in Oklahoma. We sampled invertebrates from 58 wetlands during 2009 and 2010. Diversity and taxa richness increased as the season progressed and with vegetation complexity and cover. Diversity and richness decreased as water quality was impacted by nutrient and sediment loading. Local variables occurred more consistently in taxa models than landscape variables. Important local variables included wetland hydrology, vegetation complexity, and water quality, while important landscape variables included density and type of wetlands surrounding wetlands. Land-use practice was the least important landscape variable, but is an important variable due to potential relationships with local variables such as water quality. Low variation (12–26 %) explained by the pCCA suggests other variables may be influencing invertebrate communities, but an alternative explanation is that invertebrates are insensitive to environmental variation. These findings can guide both local management of wetlands and conservation strategies at the watershed or regional scale to benefit not only invertebrates but other wetland dependent species.

The effects of different rice cultivation systems and ages on resting stages of wetland invertebrates in southern Brazil

The use of machinery and pesticides in modern agriculture are supposed to negatively affect the viability and diversity of invertebrate egg banks. Studies that evaluate the emergence of invertebrates through analysis of their dormant stages are needed in order to identify the resiliency of food webs in wetlands. In this study, the following hypotheses were tested: (1) richness is lower in conventional rice fields than in organic ones and natural wetlands; (2) richness is lower in older rice fields than younger fields; (3) composition varies among different rice systems and ages; and (4) emergence rate may vary among rice systems and ages over time. Dry sediment samples were collected from six rice fields cultivated under different systems, nine conventional rice fields with different cultivation ages and three natural wetlands. Our study showed that the richness was surprisingly higher in conventional rice fields than wetlands, with cladocerans being more associated with the former and insects more frequent in the latter. Invertebrate richness was similar among rice fields of different ages, but the composition differed between 20-year-old fields and younger fields. Variation in the viable resting stages found across different rice cultivation systems and ages contributes valuable information to wetland restoration initiatives.

Does Dispersal Limit Beetle Re‐colonization of Restored Fenland? A Case Study Using Direct Measurements of Dispersal and Genetic Analysis

AbstractDispersal ability is known to limit the colonization by many fenland species onto restored habitat but factors determining colonization by wetland invertebrates onto restored fenland have not been previously examined. We used mark‐release‐recapture and harmonic radar tracking coupled with genetic analysis (ISSR‐PCR, inter‐simple sequence repeat—polymerase chain reaction) to assess the population structure and colonization of a brachypterous wetland carabid (Carabus granulatus) onto restored fenland. The first two methods were used to determine the dispersal rate of C. granulatus in the absence of barriers, while the genetic analysis was used to examine whether populations were separated by either distance or the water channels and drainage ditches that intersect fenland landscapes. Mark‐release‐recapture and harmonic radar tracking indicated that the most active 6% of individuals could travel over 25 m in a day and individuals found to be traveling these distances remained atypically active throughout the season. Thus, C. granulatus can disperse at a rate great enough to colonize all areas of restored fenland in the study site within 1 year in the absence of barriers. We found no evidence of genetically distinct sub‐populations of C. granulatus showing that distance and water barriers did not limit the movement of individuals from potential source populations to the restored habitat. This research indicates that where fenland is created with nearby established fenland, colonization opportunities should not be considered a limiting factor to colonization by wetland carabids, even where waterways divide sites.

Amphibians in the climate vise: loss and restoration of resilience of montane wetland ecosystems in the western US

Wetlands in the remote mountains of the western US have undergone two massive ecological “experiments” spanning the 20th century. Beginning in the late 1800s and expanding after World War II, fish and wildlife managers intentionally introduced millions of predatory trout (primarily Oncorhynchus spp) into fishless mountain ponds and lakes across the western states. These new top predators, which now occupy 95% of large mountain lakes, have limited the habitat distributions of native frogs, salamanders, and wetland invertebrates to smaller, more ephemeral ponds where trout do not survive. Now a second “experiment” – anthropogenic climate change – threatens to eliminate many of these ephemeral habitats and shorten wetland hydroperiods. Caught between climate‐induced habitat loss and predation from introduced fish, native mountain lake fauna of the western US – especially amphibians – are at risk of extirpation. Targeted fish removals, guided by models of how wetlands will change under future climate scenarios, provide innovative strategies for restoring resilience of wetland ecosystems to climate change.

Evaluation of a new sampling method for assessing Cladocera richness (Crustacea, Branchiopoda) in macrophyte-rich wetlands

The wetlands found in the Brazilian Cerrado are poorly studied environments regarding ecologi- cal aspects. Assessing the diversity of aquatic invertebrates in wetlands is a challenging task, since there are no standard sampling methods that minimize the spatial effects caused by macrophytes. The aim of this study was to evaluate the efficiency of a new sampling method for assessing Cladocera richness in macrophyte-rich wetlands of Brazilian Cerrado. In six wetlands, one transect was established which corresponded to a gradient of depth or change in aquatic vegetation. Samples containing cladocerans were collected using plankton net dragged among aquatic vegetation in the dry and rainy seasons. The species accumulation curves using non-parametric estimators and the overestimation of richness were used to determine the sampling efficiency. The species accumulation curves showed different asymptotic trends regarding the season and wetland stu- died. Especially in the rainy season, an asymptotic trend was not observed in two of the wetlands studied, which may reflect the influence of seasonality on Cladocera assemblage. Nevertheless, the overestimation of species richness showed that the method of sampling was able to find more than 60% of the estimated species richness, regardless of season or wetland studied. These results indicate that the method employed for sampling Cladocera in the Cerrado wetlands can be considered adequate.

Spatial and temporal trends in invertebrate communities of Great Lakes coastal wetlands, with emphasis on Saginaw Bay of Lake Huron

The shallow-sloping coastal bathymetry of Saginaw Bay (Lake Huron) supports broad fringing wetlands. Because benthic invertebrates form an important forage base for fish, wading birds, and waterfowl that utilize these habitats, understanding the drivers of invertebrate community structure has significant management implications. We used Great Lakes basin-wide data from 2002 to place Saginaw Bay wetland invertebrate communities and their environmental drivers into a basin-wide context. Various aspects of community structure were highly correlated with fetch and watershed agriculture across the basin. Saginaw Bay wetlands had relatively high fetch and watershed agriculture and supported unique invertebrate communities, typified by high abundances of many insect taxa. Wetlands from other regions around the basin tended to have more crustaceans and gastropods than the Saginaw Bay wetlands. A 1997–2012 time series from three representative Saginaw Bay wetlands revealed substantial shifts in community structure throughout the period, especially from 2001 through 2004. These years followed a 1-m decline in Lake Huron water levels that occurred between 1997 and 2000. Major community changes included decreasing insect abundance, especially chironomids, and increasing crustacean abundances, especially Hyalella azteca (Amphipoda). While factors in addition to water levels were likely also important, our time series analysis reveals the marked temporal dynamics of Saginaw Bay wetland invertebrate communities and suggests that water level decline may have influenced these communities substantially. Both the spatial and temporal community patterns that we found should be considered in future bio-assessments utilizing wetland invertebrates.

Weak Relationships Between Environmental Factors and Invertebrate Communities in Constructed Wetlands

Growing awareness of essential wetland functions is providing support for wetland construction projects. Biomonitoring using invertebrates is a common way to evaluate project success, but relationships between wetland invertebrates and environmental factors are often weak. In recently constructed wetlands on Maryland’s Eastern Shore, we tested the hypothesis that focusing on predator and primary consumer invertebrate assemblages versus the entire community would elucidate stronger relationships with environmental factors. Despite variation in factors that are hypothesized to control wetland invertebrates (e.g., vegetation and tendency to dry), our results indicated weak relationships between environmental factors and the structure and composition of the entire community as well as predator and primary consumer assemblages separately. Examining the entire community and individual assemblages, however, showed that invertebrates were influenced by temporal factors. We propose that a complex interaction between wetland shape/size, local weather, and seasonal changes may have driven invertebrate community patterns among wetlands. Such interactions would complicate bioassessments of wetlands that differ in size, hydrology, and local weather conditions. Further study of specific factors controlling wetland invertebrates and developing new metrics that incorporate seasonal environmental change could improve biomonitoring results and thus management strategies aimed at enhancing wetland function.

Functional and Phylogenetic Relatedness in Temporary Wetland Invertebrates: Current Macroecological Patterns and Implications for Future Climatic Change Scenarios

In freshwater ecosystems, species compositions are known to be determined hierarchically by large to small‑scale environmental factors, based on the biological traits of the organisms. However, in ephemeral habitats this heuristic framework remains largely untested. Although temporary wetland faunas are constrained by a local filter (i.e., desiccation), we propose its magnitude may still depend on large-scale climate characteristics. If this is true, climate should be related to the degree of functional and taxonomic relatedness of invertebrate communities inhabiting seasonal wetlands. We tested this hypothesis in two ways. First, based on 52 biological traits for invertebrates, we conducted a case study to explore functional trends among temperate seasonal wetlands differing in the harshness (i.e., dryness) of their dry season. After finding evidence of trait filtering, we addressed whether it could be generalized across a broader climatic scale. To this end, a meta-analysis (225 seasonal wetlands spread across broad climatic categories: Arid, Temperate, and Cold) allowed us to identify whether an equivalent climate-dependent pattern of trait richness was consistent between the Nearctic and the Western Palearctic. Functional overlap of invertebrates increased from mild (i.e., Temperate) to harsher climates (i.e., Arid and Cold), and phylogenetic clustering (using taxonomy as a surrogate) was highest in Arid and lowest in Temperate wetlands. We show that, (i) as has been described in streams, higher relatedness than would be expected by chance is generally observed in seasonal wetland invertebrate communities; and (ii) this relatedness is not constant but climate-dependent, with the climate under which a given seasonal wetland is located determining the functional overlap and the phylogenetic clustering of the community. Finally, using a space-for-time substitution approach we suggest our results may anticipate how the invertebrate biodiversity embedded in these vulnerable and often overlooked ecosystems will be affected by long-term climate change.