Water beetles (Coleoptera) associated with Afrotemperate Forest patches in the Garden Route National Park, South Africa.

Southern Afrotemperate Forest is concentrated in the southern Cape region of South Africa and whilst it is relatively well known botanically, the fauna, specifically the aquatic invertebrate fauna, is poorly documented. The majority of remaining intact forest habitat is contained within the Garden Route National Park (GRNP), which straddles the provincial boundary between the Western and Eastern Cape. This study undertakes a survey of the water beetle fauna inhabiting the GRNP. The aquatic ecosystems within temperate forests of the region are poorly researched from an ecological and biodiversity perspective, despite being known to harbour endemic invertebrate elements. We collected water beetles and in situ physico-chemical data from a total of 31 waterbodies across the park over two seasons (summer and late winter) in 2017. The waterbodies sampled were mostly small freshwater perennial streams and isolated forest ponds. A total of 61 beetle taxa was recorded (29 Adephaga, 32 Polyphaga) from these waterbodies. The water beetle fauna of these forests appears to be diverse and contains many species endemic to the fynbos-dominated Cape Floristic Region, but very few of the species appear to be forest specialists. This is in contrast to the fynbos heathland habitat of the region, which harbours a high number of water beetle species endemic to this habitat, often with Gondwanan affinity. Our study is the first to document the water beetles of Afrotemperate Forests in the southern Cape region and provides an important baseline for future work on such habitats in the region and in other parts of southern Africa.

Eupotemus tuberculatus sp. nov., the first hooded shore beetle confirmed from southern Africa (Coleoptera, Epimetopidae).

Eupotemus tuberculatus sp. nov. is described on the basis of two specimens discovered during recent field work in South Africa. This is the first member of the Epimetopidae to be confirmed in Africa south of the Congo Basin and Tanzania. The new species is a member of the Eupotemus carinaticollis Fikek et al., 2021 species group, sharing features of pronotal and aedeagal design. Characters distinguishing the new species from previously described taxa are discussed and notes provided on the collecting circumstances.

Perils of life on the edge: Climatic threats to global diversity patterns of wetland macroinvertebrates.

Climate change is rapidly driving global biodiversity declines. How wetland macroinvertebrate assemblages are responding is unclear, a concern given their vital function in these ecosystems. Using a data set from 769 minimally impacted depressional wetlands across the globe (467 temporary and 302 permanent), we evaluated how temperature and precipitation (average, range, variability) affects the richness and beta diversity of 144 macroinvertebrate families. To test the effects of climatic predictors on macroinvertebrate diversity, we fitted generalized additive mixed-effects models (GAMM) for family richness and generalized dissimilarity models (GDMs) for total beta diversity. We found non-linear relationships between family richness, beta diversity, and climate. Maximum temperature was the main climatic driver of wetland macroinvertebrate richness and beta diversity, but precipitation seasonality was also important. Assemblage responses to climatic variables also depended on wetland water permanency. Permanent wetlands from warmer regions had higher family richness than temporary wetlands. Interestingly, wetlands in cooler and dry-warm regions had the lowest taxonomic richness, but both kinds of wetlands supported unique assemblages. Our study suggests that climate change will have multiple effects on wetlands and their macroinvertebrate diversity, mostly via increases in maximum temperature, but also through changes in patterns of precipitation. The most vulnerable wetlands to climate change are likely those located in warm-dry regions, where entire macroinvertebrate assemblages would be extirpated. Montane and high-latitude wetlands (i.e., cooler regions) are also vulnerable to climate change, but we do not expect entire extirpations at the family level.

Cross-continental analysis of coastal biodiversity change.

Whereas the anthropogenic impact on marine biodiversity is undebated, the quantification and prediction of this change are not trivial. Simple traditional measures of biodiversity (e.g. richness, diversity indices) do not capture the magnitude and direction of changes in species or functional composition. In this paper, we apply recently developed methods for measuring biodiversity turnover to time-series data of four broad taxonomic groups from two coastal regions: the southern North Sea (Germany) and the South African coast. Both areas share geomorphological features and ecosystem types, allowing for a critical assessment of the most informative metrics of biodiversity change across organism groups. We found little evidence for directional trends in univariate metrics of diversity for either the effective number of taxa or the amount of richness change. However, turnover in composition was high (on average nearly 30% of identities when addressing presence or absence of species) and even higher when taking the relative dominance of species into account. This turnover accumulated over time at similar rates across regions and organism groups. We conclude that biodiversity metrics responsive to turnover provide a more accurate reflection of community change relative to conventional metrics (absolute richness or relative abundance) and are spatially broadly applicable. This article is part of the theme issue 'Integrative research perspectives on marine conservation'.

Increasing salinity drastically reduces hatching success of crustaceans from depression wetlands of the semi-arid Eastern Cape Karoo region, South Africa.

Salinity is an important factor affecting freshwater aquatic species distribution and diversity. The semi-arid Eastern Cape Karoo region of South Africa has been earmarked for shale gas development through hydraulic fracturing. The process uses large amounts of water and produces briny wastewater. When not managed properly, these wastewaters may lead to salinisation of surface freshwater bodies in the region. Therefore, the effect of salinity on the hatching success of crustacean resting eggs was examined using sediments from four depression wetlands found in the region. The sediments were exposed for 28 days to salinity levels of 0.5 g L-1, 2.5 g L-1, 5 g L-1 and 10 g L-1. Control aquaria in which no salt was added were also set up. There was a significant decrease in the emerged taxa richness and abundances at salinities of 2.5 g L-1 and above. Anostraca, Notostraca and Spinicaudata hatchlings were abundant at salinities of 0.5 g L-1 and below, while Copepoda, Daphniidae (Cladocera) and Ostracoda were observed in the highest salinity, but their densities were still lower with increased salinities. Given the importance of large branchiopods in the trophic balance of depression wetlands, their loss may alter the ecological balance and function of these ecosystems.

Grazer responses to variable macroalgal resource conditions facilitate habitat structuring.

Consumer responses to altered resource conditions can vary depending on dietary preference, resource characteristics and secondary resource features such as shelter. These can have cascading effects, especially if the consumed resource impacts on overall ecological functioning. In this study, we assessed the dietary composition of grazer communities following seasonal changes in the characteristics of their staple food-source (macroalgae). This was conducted in the living stromatolite pools growing along the coast of South Africa. Stable isotope mixing models suggested that following macroalgal bleaching in summer, metazoan consumers shifted their diet from predominantly macroalgae to a generalist composition. This has important implications for the integrity of the stromatolite matrix and its layered deposition. Where previously in winter stromatolite microalgae comprised a minor component of metazoan consumer diets, in summer, following a change in the resource conditions of macroalgae, microalgae featured more prominently in grazer diets. This seasonal grazing pressure on stromatolite-related resources probably promotes the pattern of annual layering observed in the stromatolite accretion. It also demonstrates a mechanism whereby grazer dietary shifts following a change in their preferred food resource can affect the ecosystem structure of their environment, specifically the stromatolite layering process which responds to microalgal growth or grazing conditions.

Distribution and diversity of aquatic macroinvertebrate assemblages in a semi-arid region earmarked for shale gas exploration (Eastern Cape Karoo, South Africa).

This study aims to investigate macroinvertebrate assemblage structure and composition across the three major waterbody types (temporary rivers, depression wetlands and semi-permanent dams) of the Eastern Cape Karoo, and to identify important environmental and spatial correlates of macroinvertebrate assemblage composition in the region. A total of 33 waterbodies (9 dams, 13 depression wetlands and 11 rivers) were sampled. Altogether, 91 taxa were recorded in November 2014 and 82 in April 2015. Twenty-seven taxa were common to all three waterbody types (across both sampling occasions), with 17 of these observed in November and 19 in April. The ANOSIM tests revealed significant differences in assemblage composition between the depression wetlands and rivers for both sampling occasions, but dams did not differ from the other waterbody types. SIMPER analyses indicated that the notonectid Anisops varia and the corixid Micronecta scutellaris were abundant across all three waterbody types during both sampling occasions. The mayfly Cloeon africanum and the damselfly Pseudagrion sp. were abundant in river habitats during both sampling occasions, while the gastropod mollusc Bulinus tropicus and the copepod Lovenula falcifera best characterised depression wetlands on both occasions. Non-metric multidimensional scaling ordination highlighted a clear separation of assemblages between November and April, while distance-based Redundancy Analysis revealed that conductivity, altitude, turbidity and pH were the most important variables explaining the variation in macroinvertebrate assemblage patterns. These results provide baseline information which is important for future biological monitoring of impacts associated with hydraulic fracturing activities and climatic changes in the region.

Diversity and distribution of polyphagan water beetles (Coleoptera) in the Lake St Lucia system, South Africa.

Water beetles belonging to the suborder Polyphaga vary greatly in larval and adult ecologies, and fulfil important functional roles in shallow-water ecosystems by processing plant material, scavenging and through predation. This study investigates the species richness and composition of aquatic polyphagan assemblages in and around the St Lucia estuarine lake (South Africa), within the iSimangaliso Wetland Park, a UNESCO World Heritage Site. A total of 32 sites were sampled over three consecutive collection trips between 2013 and 2015. The sites encompassed a broad range of aquatic habitats, being representative of the variety of freshwater and estuarine environments present on the St Lucia coastal plain. Thirty-seven polyphagan taxa were recorded during the dedicated surveys of this study, in addition to seven species-level records from historical collections. Most beetles recorded are relatively widespread Afrotropical species and only three are endemic to South Africa. Samples were dominated by members of the Hydrophilidae (27 taxa), one of which was new to science (Hydrobiomorpha perissinottoi Bilton, 2016). Despite the fauna being dominated by relatively widespread taxa, five represent new records for South Africa, highlighting the poor state of knowledge on water beetle distribution patterns in the region. Wetlands within the dense woodland characterising the False Bay region of St Lucia supported a distinct assemblage of polyphagan beetles, whilst sites occurring on the Eastern and Western Shores of Lake St Lucia were very similar in their beetle composition. In line with the Afrotropical region as a whole, the aquatic Polyphaga of St Lucia appear to be less diverse than the Hydradephaga, for which 68 species were recorded during the same period. However, the results of the present study, in conjunction with those for Hydradephaga, show that the iSimangaliso Wetland Park contains a high beetle diversity. The ongoing and future ecological protection of not only the estuarine lake itself, but also surrounding freshwater wetlands, is imperative and should be taken into consideration during future management planning for the park.

Non-reliance of metazoans on stromatolite-forming microbial mats as a food resource.

Grazing and burrowing organisms usually homogenise microalgal mats that form on benthic sediments of many aquatic ecosystems. In the absence of this disruption, microalgal mats can accrete laminated deposits (stromatolites). Stromatolites are rare in modern coastal ecosystems, but persist at locations where metazoans are largely excluded. This study aimed to assess the trophic structure at stromatolite locations where metazoans co-occur, to determine the grazing influence exerted by the metazoans on the stromatolite-forming microalgae (cyanobacteria and diatoms). Stable isotope signatures (δ13C and δ15N) were used as food-web tracers and dietary composition of consumers was calculated using source mixing models. Results clearly demonstrate that the dominant macrofaunal grazers do not utilise stromatolite material as a food resource, but rather subsist on autochthonous macroalgae. For instance, the mean (±SD) dietary composition of two of the most abundant grazers, Melita zeylanica (Amphipoda) and Composetia cf. keiskama (Polychaeta), consisted of 80 ± 11% and 91 ± 7% macroalgae, respectively. This suggests that the stromatolite-forming benthic microalgae are not disrupted significantly by grazing pressures, allowing for the layered mineralisation process to perpetuate. Additionally, grazers likely have a restrictive influence on pool macroalgae, maintaining the competitive balance between micro- and macroalgal groups.

Ecology and distribution of large branchiopods (Crustacea, Branchiopoda, Anostraca, Notostraca, Laevicaudata, Spinicaudata) of the Eastern Cape Karoo, South Africa.

A survey of the large branchiopod fauna of the Eastern Cape Karoo region of South Africa was undertaken to provide baseline biodiversity information in light of impending shale gas development activities in the region. Twenty-two waterbodies, including nine dams and thirteen natural depression wetlands, were sampled during November 2014 and April 2015. A total of 13 species belonging to four orders were collected, comprising five anostracans, one notostracan, six spinicaudatans and one laevicaudatan. Cyzicus australis was most common, occurring in 46% of the waterbodies. Species co-occurred in 87% of the waterbodies, with a maximum number of six species recorded from the same waterbody. Our new distribution records for Lynceus truncatus, Streptocephalus spinicaudatus and Streptocephalus indistinctus represent substantial expansions of the previously known ranges for these species. Tarkastad is now the westernmost record for Streptocephalus spinicaudatus, while Jansenville now constitutes the southernmost record for Streptocephalus indistinctus. Large branchiopod distribution data from previous Eastern Cape records were combined with our current data, demonstrating that a total of 23 large branchiopod species have been recorded from the region to date. As the Karoo is one of the few major shale basins in the world where the natural baseline is still largely intact, this survey forms a basis for future reference and surface water quality monitoring during the process of shale gas exploration/extraction.

Predaceous water beetles (Coleoptera, Hydradephaga) of the Lake St Lucia system, South Africa: biodiversity, community ecology and conservation implications.

Water beetles are one of the dominant macroinvertebrate groups in inland waters and are excellent ecological indicators, reflecting both the diversity and composition of the wider aquatic community. The predaceous water beetles (Hydradephaga) make up around one-third of known aquatic Coleoptera and, as predators, are a key group in the functioning of many aquatic habitats. Despite being relatively well-known taxonomically, ecological studies of these insects in tropical and subtropical systems remain rare. A dedicated survey of the hydradephagan beetles of the Lake St Lucia wetlands (South Africa) was undertaken between 2013 and 2015, providing the first biodiversity census for this important aquatic group in the iSimangaliso Wetland Park, a UNESCO World Heritage Site within the Maputaland biodiversity hotspot. A total of 32 sites covering the entire spectrum of waterbody types were sampled over the course of three collecting trips. The Lake St Lucia wetlands support at least 68 species of Hydradephaga, a very high level of diversity comparing favourably with other hotspots on the African continent and elsewhere in the world and a number of taxa are reported for South Africa for the first time. This beetle assemblage is dominated by relatively widespread Afrotropical taxa, with few locally endemic species, supporting earlier observations that hotspots of species richness and centres of endemism are not always coincident. Although there was no significant difference in the number of species supported by the various waterbody types sampled, sites with the highest species richness were mostly temporary depression wetlands. This contrasts markedly with the distribution of other taxa in the same system, such as molluscs and dragonflies, which are most diverse in permanent waters. Our study is the first to highlight the importance of temporary depression wetlands and emphasises the need to maintain a variety of wetland habitats for aquatic conservation in this biodiverse region.

Coexisting living stromatolites and infaunal metazoans.

Microbialites, bioaccretionary structures formed during the growth and metabolism of microorganisms (principally cyanobacteria) were the dominant lifeform in shallow late-Archean and Proterozoic oceans. During the Cambrian radiation of metazoan life, which began ~540 Mya, microbialite abundance and diversity further declined following a peak in the Mesoproterozoic. Notwithstanding contention, grazing and bioturbation effects of metazoans have been hypothesized as the dominant driver of modern microbialite scarcity. However, this metazoan-microbialite exclusion has not been fully explored in the few extant microbialites. Here we provide further evidence showing that living marine layered microbialites (stromatolites) coexist with a persistent assemblage of benthic macro-invertebrates, as has previously been demonstrated in some thrombolitic (clotted) microbialites. Surprisingly, these metazoans have active habits, such as burrowing, which should be expected to disrupt the layered matrix. As other studies have shown, through a network of burrows, metazoans can exploit local diurnal oxygen refugia within microbialites as well as escape predation. Our results, therefore, add novel evidence in support of the hypotheses that geologically, metazoans are not always incompatible with stromatolites, while ecologically, microbialites may act as micro-refugia for modern metazoans and historically have performed a similar inferred role in past ecosystems.

Wetlands in changed landscapes: the influence of habitat transformation on the physico-chemistry of temporary depression wetlands.

Temporary wetlands dominate the wet season landscape of temperate, semi-arid and arid regions, yet, other than their direct loss to development and agriculture, little information exists on how remaining wetlands have been altered by anthropogenic conversion of surrounding landscapes. This study investigates relationships between the extent and type of habitat transformation around temporary wetlands and their water column physico-chemical characteristics. A set of 90 isolated depression wetlands (seasonally inundated) occurring on coastal plains of the south-western Cape mediterranean-climate region of South Africa was sampled during the winter/spring wet season of 2007. Wetlands were sampled across habitat transformation gradients according to the areal cover of agriculture, urban development and alien invasive vegetation within 100 and 500 m radii of each wetland edge. We hypothesized that the principal drivers of physico-chemical conditions in these wetlands (e.g. soil properties, basin morphology) are altered by habitat transformation. Multivariate multiple regression analyses (distance-based Redundancy Analysis) indicated significant associations between wetland physico-chemistry and habitat transformation (overall transformation within 100 and 500 m, alien vegetation cover within 100 and 500 m, urban cover within 100 m); although for significant regressions the amount of variation explained was very low (range: ∼2 to ∼5.5%), relative to that explained by purely spatio-temporal factors (range: ∼35.5 to ∼43%). The nature of the relationships between each type of transformation in the landscape and individual physico-chemical variables in wetlands were further explored with univariate multiple regressions. Results suggest that conservation of relatively narrow (∼100 m) buffer strips around temporary wetlands is likely to be effective in the maintenance of natural conditions in terms of physico-chemical water quality.