Spatial versus spatio-temporal approaches for studying metacommunities: a multi-taxon analysis in Mediterranean and tropical temporary ponds.

Prior research on metacommunities has largely focused on snapshot surveys, often overlooking temporal dynamics. In this study, our aim was to compare the insights obtained from metacommunity analyses based on a spatial approach repeated over time, with a spatio-temporal approach that consolidates all data into a single model. We empirically assessed the influence of temporal variation in the environment and spatial connectivity on the structure of metacommunities in tropical and Mediterranean temporary ponds. Employing a standardized methodology across both regions, we surveyed multiple freshwater taxa in three time periods within the same hydrological year from multiple temporary ponds in each region. To evaluate how environmental, spatial and temporal influences vary between the two approaches, we used nonlinear variation partitioning analyses based on generalized additive models. Overall, this study underscores the importance of adopting spatio-temporal analytics to better understand the processes shaping metacommunities. While the spatial approach suggested that environmental factors had a greater influence, our spatio-temporal analysis revealed that spatial connectivity was the primary driver influencing metacommunity structure in both regions. Temporal effects were equally important as environmental effects, suggesting a significant role of ecological succession in metacommunity structure.

Integrative Taxonomy of Cyclocyprididae Kaufmann, 1900 (Ostracoda: Podocopa) with Description of a New Genus and Species.

The two widespread ostracod genera Cypria Zenker, 1854 and Physocypria Vávra, 1897 are traditionally distinguished based on the presence or absence of tubercles on the right valve margin. However, recent research based on soft body parts has uncovered new cryptic genera within Cypria and Physocypria. Following this line of research, a new Cyclocyprididae genus and species, Vizcainocypria viator gen. nov. sp. nov., is here described from individuals collected in rice fields and wetlands of the Iberian Peninsula. Vizcainocypria is compared with Cypria, Physocypria, Dentocypria Savatenalinton, 2017, Keysercypria Karanovic, 2011, Brasilocypria Almeida et al., 2023, and Claudecypria Almeida et al., 2023 based on morphological evidence. Besides the presence or absence of tubercles on the right valve, these genera can be distinguished according to their mandibular palp, second thoracopod, caudal ramus, and male hemipenis. Molecular analyses using mitochondrial (COX1), and nuclear (28S rDNA) genes provide further support for the differentiation of Cypria, Dentocypria, Physocypria and Vizcainocypria gen. nov. The present study highlights the importance of using an integrative taxonomy approach, combining shell and soft-body parts morphology and molecular data, to characterize the rich diversity of freshwater ostracods.

Understanding Ecological Complexity in a Chemical Stress Context: A Reflection on Recolonization, Recovery, and Adaptation of Aquatic Populations and Communities.

Recovery, recolonization, and adaptation in a chemical stress context are processes that regenerate local populations and communities as well as the functions these communities perform. Recolonization, either by species previously present or by new species able to occupy the niches left empty, refers to a metacommunity process with stressed ecosystems benefiting from the dispersal of organisms from other areas. A potential consequence of recolonization is a limited capacity of local populations to adapt to potentially repeating events of chemical stress exposure when their niches have been effectively occupied by the new colonizers or by new genetic lineages of the taxa previously present. Recovery, instead, is an internal process occurring within stressed ecosystems. More specifically, the impact of a stressor on a community benefits less sensitive individuals of a local population as well as less sensitive taxa within a community. Finally, adaptation refers to phenotypic and, sometimes, genetic changes at the individual and population levels, allowing the permanence of individuals of previously existing taxa without necessarily changing the community taxonomic composition (i.e., not replacing sensitive species). Because these processes are usually operating in parallel in nature, though at different degrees, it seems relevant to try to understand their relative importance for the regeneration of community structure and ecosystem functioning after chemical exposure. In the present critical perspective, we employed case studies supporting our understanding of the underlying processes with the hope to provide a theoretical framework to disentangle the relevance of the three processes for the regeneration of a biological community after chemical exposure. Finally, we provide some recommendations to experimentally compare their relative importance so that the net effects of these processes can be used to parameterize risk-assessment models and inform ecosystem management. Environ Toxicol Chem 2023;42:1857-1866. © 2023 The Authors. Environmental Toxicology and Chemistry published by Wiley Periodicals LLC on behalf of SETAC.

Socio-ecological impact of monogenetic volcanism in the La Garrotxa Volcanic Field (NE Iberia).

Volcanism can cause major impacts, including climate change and mass extinctions. However, the impact of monogenetic volcanism is often considered as limited in volcanological research. This work provides for the first time an interdisciplinary approach to the socio-ecological impact of monogenetic volcanism in a key region, the La Garrotxa Volcanic Field (GVF, Girona, NE Iberia), where intense monogenetic volcanic activity occurred in the past. The analyses of a sedimentary sequence from the GVF enabled identifying previously unknown volcanic eruptions in the time interval 14-8.4 ka cal BP, constrain their volcanic stratigraphy and age, and unfold the effects of environmental change on geomorphology, vegetation, aquatic organisms and humans. Moreover, we reconstruct the major palaeoenvironmental changes caused by the eruptions in terms of fire episodes and subsequent disturbance on vegetation, hydrology and limnological conditions. When put in context with the archaeological record, it appears that the last hunter-gatherer communities were resilient at an extra-local scale, facing episodes of vulnerability due to volcanic activity, suggesting that their flexible nomadic patterns and foraging economies were an efficient source of risk management against the volcanic eruptions and their ecological impacts.

Inconsistent response of taxonomic groups to space and environment in mediterranean and tropical pond metacommunities.

The metacommunity concept provides a theoretical framework that aims at explaining organism distributions by a combination of environmental filtering, dispersal, and drift. However, few works have attempted a multitaxon approach and even fewer have compared two distant biogeographical regions using the same methodology. We tested the expectation that temperate (mediterranean-climate) pond metacommunities would be more influenced by environmental and spatial processes than tropical ones, because of stronger environmental gradients and a greater isolation of waterbodies. However, the pattern should be different among groups of organisms depending on their dispersal abilities. We surveyed 30 tropical and 32 mediterranean temporary ponds from Costa Rica and Spain, respectively, and obtained data on 49 environmental variables. We characterized the biological communities of bacteria and archaea (from the water column and the sediments), phytoplankton, zooplankton, benthic invertebrates, amphibians and birds, and estimated the relative role of space and environment on metacommunity organization for each group and region, by means of variation partitioning using generalized additive models. Purely environmental effects were important in both tropical and mediterranean ponds, but stronger in the latter, probably due to their larger limnological heterogeneity. Spatially correlated environment and pure spatial effects were greater in the tropics, related to higher climatic heterogeneity and dispersal processes (e.g., restriction, surplus) acting at different scales. The variability between taxonomic groups in the contribution of spatial and environmental factors to metacommunity variation was very wide, but higher in active, compared with passive, dispersers. Higher environmental effects were observed in mediterranean passive dispersers, and higher spatial effects in tropical passive dispersers. The unexplained variation was larger in the tropical setting, suggesting a higher role for stochastic processes, unmeasured environmental factors, or biotic interactions in the tropics, although this difference affected some actively dispersing groups (insects and birds) more than passive dispersers. These results, despite our limitations in comparing only two regions, provide support, for a wide variety of aquatic organisms, for the classic view of stronger abiotic niche constraints in temperate areas compared with the tropics. The heterogeneous response of taxonomic groups between regions also points to a stronger influence of regional context than organism adaptations on metacommunity organization.

Disturbance-induced emigration: an overlooked mechanism that reduces metapopulation extinction risk.

Emigration propensity (i.e., the tendency to leave undisturbed patches) is a key life-history trait of organisms in metapopulations with local extinctions and colonizations. Metapopulation models of dispersal evolution typically assume that patch disturbance kills all individuals within the patch, thus causing local extinction. However, individuals may instead be able to leave a patch when it is disturbed, either by fleeing before being killed or simply because the disturbance destroys the patch without causing mortality. This scenario may pertain to a wide range of organisms from horizontally transmitted symbionts, to aquatic insects inhabiting temporary ponds, to vertebrates living in fragmented forests. We generalized a Levins-type metapopulation model of dispersal evolution by adding a new parameter of disturbance escape probability, which incorporates a second source of dispersal into the model: disturbance-induced emigration. We show that disturbance escape expands the domain of metapopulation viability and selects for lower rates of emigration propensity when disturbance rates are high. The fitness gains from disturbance-induced emigration are generally moderate, suggesting that disturbance escape might act more as a complementary dispersal strategy rather than a replacement to emigration propensity, at least for metapopulations that meet the assumptions of the Levins-type model. Yet disturbance-induced emigration may in some circumstances rescue a metapopulation from long-term extinction when the combination of high disturbance rates and low local population growth rates compromises its viability. Further, a metapopulation could persist exclusively by disturbance escape if local carrying capacities are large enough to counterbalance two sources of mortality: mortality driven by disturbance and mortality during dispersal. This study opens two promising research lines: (1) the investigation of disturbance escape in metapopulations of ephemeral habitats with unsaturated populations and non-equilibrium dynamics and (2) the incorporation of information costs to investigate the joint evolution of disturbance escape and emigration propensity.

A new species of Cypris (Crustacea: Ostracoda) from the Iberian Peninsula and the Balearic Islands, with comments on the first ostracod named using the Linnean system.

The genus Cypris, considered the oldest ostracod generic name erected using the Linnean system, comprises a reduced number of large-bodied species, mostly found in Africa and Asia. Only six of them are known to occur in Europe. Here we describe a new species, Cypris pretusi sp. nov., collected in small temporary streams and ponds along the Eastern Iberian Peninsula and Minorca (Balearic Islands). The new species is very close to the type species of the genus, Cypris pubera O.F. Müller, 1776, but differs from it in having a set of smaller subequal spines on the posterior edge of the valves, by the absence of conspicuous spines along the front edge, and by the beak-like frontal shape of its carapace in dorsal view, similar to Cypris decaryi Gauthier, 1933. Soft parts are very similar to the type species, but it differs in having shorter swimming setae on the second antennae. Molecular analyses of the COX1 region support its status as a species distinct from C. pubera and closer to Cypris bispinosa Lucas, 1849, also providing evidence for a separation of C. pubera s.l. in two clades, one of which is here considered to correspond to Cypris triaculeata Daday, 1892. We discuss the relationships of C. pretusi sp. nov. to other members of the genus and its possible origin from nearby biogeographic regions (probably Africa or Asia) and provide a key to species of Cypris found in Europe. We also discuss the relationship between Monoculus concha pedata (= M. conchaceus), the first ostracod named by Linnæus, and Cypris pubera, the type species of the genus, described by Müller in 1776 and considered by him the same species as the one first named by Linnæus.

The interplay of nested biotic interactions and the abiotic environment regulates populations of a hypersymbiont.

The role of biotic interactions in shaping the distribution and abundance of species should be particularly pronounced in symbionts. Indeed, symbionts have a dual niche composed of traits of their individual hosts and the abiotic environment external to the host, and often combine active dispersal at finer scales with host-mediated dispersal at broader scales. The biotic complexity in the determinants of species distribution and abundance should be even more pronounced for hyper symbionts (symbionts of other symbionts). We use a chain of symbiosis to explore the relative influence of nested biotic interactions and the abiotic environment on occupancy and abundance of a hypersymbiont. Our empirical system is the epibiont ciliate Lagenophrys discoidea, which attaches to an ostracod that is itself ectosymbiotic on crayfish (the basal host). We applied multimodel selection and variance partitioning for GLMM to assess the relative importance of (a) traits of symbiotic hosts (ostracod sex and abundance), (b) traits of basal hosts (crayfish body weight, abundance and intermoult stage), (c) the abiotic environment (water chemistry and climate) and (d) geospatial autocorrelation patterns (capturing potential effects of crayfish dispersal among localities). Our models explained about half of the variation in prevalence and abundance of the hypersymbiont. Variation in prevalence was partly explained, in decreasing order of importance (18%-4%) by shared effects of symbiotic host traits and the abiotic environment, pure fixed effects of symbiotic hosts, abiotic environment and geospatial patterns (traits of basal hosts were not relevant). Hypersymbiont abundance was most strongly explained by random effects of host traits (mainly the symbiotic host), in addition to weaker fixed effects (mostly abiotic environment). Our results highlight the major role of the interplay between abundance of symbiotic hosts and water physico-chemistry in regulating populations of a hypersymbiotic ciliate, which is likely critical for dispersal dynamics, availability of attachment resources and suitability of on-host living conditions for the ciliate. We also found moderate signal of regulation by the basal host, for which we propose three mechanisms: (a) modulation of microhabitat suitability (crayfish-created water currents); (b) concentration of symbiotic hosts within crayfish; and (c) dispersal mediated by crayfish.

Coastal Evolution in a Mediterranean Microtidal Zone: Mid to Late Holocene Natural Dynamics and Human Management of the Castelló Lagoon, NE Spain.

We present a palaeoenvironmental study of the Castelló lagoon (NE Spain), an important archive for understanding long-term interactions between dynamic littoral ecosystems and human management. Combining geochemistry, mineralogy, ostracods, diatoms, pollen, non-pollen palynomorphs, charcoal and archaeo-historical datasets we reconstruct: 1) the transition of the lagoon from a marine to a marginal environment between ~3150 cal BC to the 17th century AD; 2) fluctuations in salinity; and 3) natural and anthropogenic forces contributing to these changes. From the Late Neolithic to the Medieval period the lagoon ecosystem was driven by changing marine influence and the land was mainly exploited for grazing, with little evidence for impact on the natural woodland. Land-use exploitation adapted to natural coastal dynamics, with maximum marine flooding hampering agropastoral activities between ~1550 and ~150 cal BC. In contrast, societies actively controlled the lagoon dynamics and become a major agent of landscape transformation after the Medieval period. The removal of littoral woodlands after the 8th century was followed by the expansion of agrarian and industrial activities. Regional mining and smelting activities polluted the lagoon with heavy metals from the ~11th century onwards. The expansion of the milling industry and of agricultural lands led to the channelization of the river Muga into the lagoon after ~1250 cal AD. This caused its transformation into a freshwater lake, increased nutrient load, and the infilling and drainage of a great part of the lagoon. By tracking the shift towards an anthropogenically-controlled system around ~750 yr ago, this study points out Mediterranean lagoons as ancient and heavily-modified systems, with anthropogenic impacts and controls covering multi-centennial and even millennial timescales. Finally, we contributed to the future construction of reliable seashell-based chronologies in NE Spain by calibrating the Banyuls-sur-Mer ΔR offset with ceramic imports from the Emporiae archaeological site.

Hydrology Affects Environmental and Spatial Structuring of Microalgal Metacommunities in Tropical Pacific Coast Wetlands.

The alternating climate between wet and dry periods has important effects on the hydrology and therefore on niche-based processes of water bodies in tropical areas. Additionally, assemblages of microorganism can show spatial patterns, in the form of a distance decay relationship due to their size or life form. We aimed to test spatial and environmental effects, modulated by a seasonal flooding climatic pattern, on the distribution of microalgae in 30 wetlands of a tropical dry forest region: the Pacific coast of Costa Rica and Nicaragua. Three surveys were conducted corresponding to the beginning, the highest peak, and the end of the hydrological year during the wet season, and species abundance and composition of planktonic and benthic microalgae was determined. Variation partitioning analysis (as explained by spatial distance or environmental factors) was applied to each seasonal dataset by means of partial redundancy analysis. Our results show that microalgal assemblages were structured by spatial and environmental factors depending on the hydrological period of the year. At the onset of hydroperiod and during flooding, neutral effects dominated community dynamics, but niche-based local effects resulted in more structured algal communities at the final periods of desiccating water bodies. Results suggest that climate-mediated effects on hydrology can influence the relative role of spatial and environmental factors on metacommunities of microalgae. Such variability needs to be accounted in order to describe accurately community dynamics in tropical coastal wetlands.

Human-mediated dispersal of aquatic invertebrates with waterproof footwear.

Human-mediated dispersal has rarely been considered in wetland conservation strategies at regional scales, yet high concern exists about this aspect for (inter-)national management considering invasive species in other aquatic systems. In this context, we aim at understanding the role of human-mediated dispersal by footwear in protected wetlands with high conservation value. Zooplankton and zoobenthos were sampled in 13 shallow lakes in central Spain and, at the same time, mud attached to waders was collected and later cultured in deionized water under laboratory conditions for 4 weeks. Two-hundred and four individuals belonging to 19 invertebrate taxa were recovered after hatching; Ostracoda (84 %), Cladocera (53 %), Copepoda (30 %), Anostraca (30 %), and Notostraca (7 %) were the most frequent groups among the hatched crustaceans. NMDS and PERMANOVA analyses showed significant differences between the dispersed (via footwear) and the source active metacommunity, suggesting different dispersal abilities among the species found. Human vectors facilitate dispersal among protected lakes, which could eventuality lead to biotic homogenization and faster spread of non-indigenous species. Preservation strategies and education campaigns associated to target humans in close contact with water bodies should be imperative in conservation management of protected lakes.

Breaking the temperature-size rule: thermal effects on growth, development and fecundity of a crustacean from temporary waters.

The temperature-size rule (TSR) is a well-established phenomenon to describe the growth response of ectotherms to temperature by which individuals maintained at low temperatures grow more slowly, but attain a larger size upon maturity. Although there are adaptive and non-adaptive theories about the plasticity of body size in response to temperature, these cannot be applied to all ectotherms, and little is known about the changes in growth and development rates through ontogeny. The ostracod species Heterocypris bosniaca, an inhabitant of freshwater temporary ponds, was used to examine the growth and development rates of its nine growth stages and female fecundity at four different temperatures (15°C, 20°C, 25°C and 30°C). The development rate of this species accelerates with increasing temperature, reaching a maximum value at 25°C. The growth factor has a reverse-TSR in younger instars, and the typical TSR is followed only in the last two moults, resulting in non-monotonic response of adult size to temperature. Fecundity (total offspring per female) was not directly related to adult size and was generally higher at lower temperatures. Our results agree with recent research showing that the TSR may vary during ontogeny, and may not be a general trend in ostracod species from temporary waters. Indeed, adult carapace size seems to follow the pattern of a thermal reaction norm, probably influenced by the reduction of oxygen bioavailability at low temperature and the drastic increase in metabolic demand at the upper extreme of the thermal gradient.

Invasion biology in non-free-living species: interactions between abiotic (climatic) and biotic (host availability) factors in geographical space in crayfish commensals (Ostracoda, Entocytheridae).

In invasion processes, both abiotic and biotic factors are considered essential, but the latter are usually disregarded when modeling the potential spread of exotic species. In the framework of set theory, interactions between biotic (B), abiotic (A), and movement-related (M) factors in the geographical space can be hypothesized with BAM diagrams and tested using ecological niche models (ENMs) to estimate A and B areas. The main aim of our survey was to evaluate the interactions between abiotic (climatic) and biotic (host availability) factors in geographical space for exotic symbionts (i.e., non-free-living species), using ENM techniques combined with a BAM framework and using exotic Entocytheridae (Ostracoda) found in Europe as model organisms. We carried out an extensive survey to evaluate the distribution of entocytherids hosted by crayfish in Europe by checking 94 European localities and 12 crayfish species. Both exotic entocytherid species found, Ankylocythere sinuosa and Uncinocythere occidentalis, were widely distributed in W Europe living on the exotic crayfish species Procambarus clarkii and Pacifastacus leniusculus, respectively. No entocytherids were observed in the remaining crayfish species. The suitable area for A. sinuosa was mainly restricted by its own limitations to minimum temperatures in W and N Europe and precipitation seasonality in circum-Mediterranean areas. Uncinocythere occidentalis was mostly restricted by host availability in circum-Mediterranean regions due to limitations of P. leniusculus to higher precipitation seasonality and maximum temperatures. The combination of ENMs with set theory allows studying the invasive biology of symbionts and provides clues about biogeographic barriers due to abiotic or biotic factors limiting the expansion of the symbiont in different regions of the invasive range. The relative importance of abiotic and biotic factors on geographical space can then be assessed and applied in conservation plans. This approach can also be implemented in other systems where the target species is closely interacting with other taxa.

Different scales of spatial segregation of two species of feather mites on the wings of a passerine bird.

The "condition-specific competition hypothesis" proposes that coexistence of 2 species is possible when spatial or temporal variations in environmental conditions exist and each species responds differently to those conditions. The distribution of different species of feather mites on their hosts is known to be affected by intrinsic host factors such as structure of feathers and friction among feathers during flight, but there is also evidence that external factors such as humidity and temperature can affect mite distribution. Some feather mites have the capacity to move through the plumage rather rapidly, and within-host variation in intensity of sunlight could be one of the cues involved in these active displacements. We analyzed both the within- and between-feather spatial distribution of 2 mite species, Trouessartia bifurcata and Dolichodectes edwardsi , that coexist in flight feathers of the moustached warbler Acrocephalus melanopogon. A complex spatial segregation between the 2 species was observed at 3 spatial levels, i.e., "feather surfaces," "between feathers," and "within feathers." Despite certain overlapping distribution among feathers, T. bifurcata dominated proximal and medial regions on dorsal faces, while D. edwardsi preferred disto-ventral feather areas. An experiment to check the behavioral response of T. bifurcata to sunlight showed that mites responded to light exposure by approaching the feather bases and even leaving its dorsal face. Spatial heterogeneity across the 3 analyzed levels, together with response to light and other particular species adaptations, may have played a role in the coexistence and segregation of feather mites competing for space and food in passerine birds.