Global COVID-19 lockdown highlights humans as both threats and custodians of the environment.

The global lockdown to mitigate COVID-19 pandemic health risks has altered human interactions with nature. Here, we report immediate impacts of changes in human activities on wildlife and environmental threats during the early lockdown months of 2020, based on 877 qualitative reports and 332 quantitative assessments from 89 different studies. Hundreds of reports of unusual species observations from around the world suggest that animals quickly responded to the reductions in human presence. However, negative effects of lockdown on conservation also emerged, as confinement resulted in some park officials being unable to perform conservation, restoration and enforcement tasks, resulting in local increases in illegal activities such as hunting. Overall, there is a complex mixture of positive and negative effects of the pandemic lockdown on nature, all of which have the potential to lead to cascading responses which in turn impact wildlife and nature conservation. While the net effect of the lockdown will need to be assessed over years as data becomes available and persistent effects emerge, immediate responses were detected across the world. Thus, initial qualitative and quantitative data arising from this serendipitous global quasi-experimental perturbation highlights the dual role that humans play in threatening and protecting species and ecosystems. Pathways to favorably tilt this delicate balance include reducing impacts and increasing conservation effectiveness.

Temporal clustering of extreme climate events drives a regime shift in rocky intertidal biofilms.

Research on regime shifts has focused primarily on how changes in the intensity and duration of press disturbances precipitate natural systems into undesirable, alternative states. By contrast, the role of recurrent pulse perturbations, such as extreme climatic events, has been largely neglected, hindering our understanding of how historical processes regulate the onset of a regime shift. We performed field manipulations to evaluate whether combinations of extreme events of temperature and sediment deposition that differed in their degree of temporal clustering generated alternative states in rocky intertidal epilithic microphytobenthos (biofilms) on rocky shores. The likelihood of biofilms to shift from a vegetated to a bare state depended on the degree of temporal clustering of events, with biofilm biomass showing both states under a regime of non-clustered (60 d apart) perturbations while collapsing in the clustered (15 d apart) scenario. Our results indicate that time since the last perturbation can be an important predictor of collapse in systems exhibiting alternative states and that consideration of historical effects in studies of regime shifts may largely improve our understanding of ecosystem dynamics under climate change.

Hybrid datasets: integrating observations with experiments in the era of macroecology and big data.

Understanding how increasing human domination of the biosphere affects life on earth is a critical research challenge. This task is facilitated by the increasing availability of open-source data repositories, which allow ecologists to address scientific questions at unprecedented spatial and temporal scales. Large datasets are mostly observational, so they may have limited ability to uncover causal relations among variables. Experiments are better suited at attributing causation, but they are often limited in scope. We propose hybrid datasets, resulting from the integration of observational with experimental data, as an approach to leverage the scope and ability to attribute causality in ecological studies. We show how the analysis of hybrid datasets with emerging techniques in time series analysis (Convergent Cross-mapping) and macroecology (Joint Species Distribution Models) can generate novel insights into causal effects of abiotic and biotic processes that would be difficult to achieve otherwise. We illustrate these principles with two case studies in marine ecosystems and discuss the potential to generalize across environments, species and ecological processes. If used wisely, the analysis of hybrid datasets may become the standard approach for research goals that seek causal explanations for large-scale ecological phenomena.

Legacy effects and memory loss: how contingencies moderate the response of rocky intertidal biofilms to present and past extreme events.

Understanding how historical processes modulate the response of ecosystems to perturbations is becoming increasingly important. In contrast to the growing interest in projecting biodiversity and ecosystem functioning under future climate scenarios, how legacy effects originating from historical conditions drive change in ecosystems remains largely unexplored. Using experiments in combination with stochastic antecedent modelling, we evaluated how extreme warming, sediment deposition and grazing events modulated the ecological memory of rocky intertidal epilithic microphytobenthos (EMPB). We found memory effects in the non-clustered scenario of disturbance (60 days apart), where EMPB biomass fluctuated in time, but not under clustered disturbances (15 days apart), where EMPB biomass was consistently low. A massive grazing event impacted on EMPB biomass in a second run of the experiment, also muting ecological memory. Our results provide empirical support to the theoretical expectation that stochastic fluctuations promote ecological memory, but also show that contingencies may lead to memory loss.

Spatial signatures of an approaching regime shift in Posidonia oceanica meadows.

Determining the proximity of ecosystems to tipping points is a critical yet complex task, heightened by the growing severity of climate change and local anthropogenic stressors on ecosystem integrity. Spatial Early Warning Signals (EWS) have been recognized for their potential in preemptively signaling regime shifts to degraded states, but their performance in natural systems remains uncertain. In this study, we investigated the performance of 'recovery length' - the spatial extent of recovery from a perturbation - and spatial EWS as early warnings of regime shifts in Posidonia oceanica meadows. Our experimental approach involved progressively thinning the P. oceanica canopy, from 0 to 100%, at the edge of a dead-matte area - a structure formed by dead P. oceanica rhizomes and colonized by algal turfs - to promote the propagation of algal turfs. We calculated recovery length as the distance from the dead-matte edge to the point where algal turfs colonized the canopy-thinned region. Our results showed a linear increase in recovery length with canopy thinning, successfully anticipated the degradation of P. oceanica. While spatial skewness decline with increased canopy degradation, other spatial EWS, such as Moran correlation at lag-1, low-frequency spatial spectra, and spatial variance, were ineffective in signaling this degradation. These findings underscore the potential of recovery length as a reliable early warning indicator of regime shifts in marine coastal ecosystems.

Stabilizing effects of spatially heterogeneous disturbance via reduced spatial synchrony on a rocky shore community.

Understanding how synchronous species fluctuations affect community stability is a main research topic in ecology. Yet experimental studies evaluating how changes in disturbance regimes affect the synchrony and stability of populations and communities remain rare. We hypothesized that spatially heterogeneous disturbances of moderate intensity would promote metacommunity stability by decreasing the spatial synchrony of species fluctuations. To test this hypothesis, we exposed rocky shore communities of algae and invertebrates to homogeneous and gradient-like spatial patterns of disturbance at two levels of intensity for 4 years and used synchrony networks to characterize community responses to these disturbances. The gradient-like disturbance at low intensity enhanced spatial ß diversity compared to the other treatments and produced the most heterogeneous and least synchronized network, which was also the most stable in terms of population and community fluctuations. In contrast, homogeneous disturbance destabilized the community, enhancing spatial synchronization. Intense disturbances always reduced spatial ß diversity, indicating that strong perturbations could destabilize communities via biotic homogenization regardless of their spatial structure. Our findings corroborated theoretical predictions, emphasizing the importance of spatially heterogeneous disturbances in promoting stability by amplifying asynchronous spatial and temporal fluctuations in population and community abundance. In contrast to other networks, synchrony networks are vulnerable to the removal of most peripheral nodes, which are less synchronized, but may contribute more to stability than other nodes by dampening large fluctuations in species abundance. Our findings suggest that climate change and direct anthropogenic disturbance can compromise the stability of ecological communities through combined effects on diversity and synchrony, as well as further affecting ecosystems through habitat loss.

Assessing the risk of oil spill impacts and potential biodiversity loss for coastal marine environment at the turn of the COVID-19 pandemic event.

The Tuscan Archipelago, with its great environmental and economic importance, is one of the highest oil spill density areas in the Western Mediterranean. In this study, an interdisciplinary approach, based on numerical applications and experimental methods, was implemented to quantify the risk of oil spill impact along the rocky shores of this archipelago in relation to the maritime activities. The risk, defined as a combination of the hazard and the damage, was quantified for the biennial 2019-2020 in order to account for the effects generated by the COVID-19 pandemic restrictions on the local maritime traffic. A high-resolution oceanographic and particle tracking model was applied to simulate the trajectories of possible oil spill events and to quantify the hazard of impacts on the coast of numerical particles, daily seeded in correspondence of those marine sectors that are characterised by relevant traffic of vessels. The damage, expressed as the product of exposure and vulnerability, was estimated following an extensive sampling approach aimed at quantifying the ecological status of the rocky shores in four selected islands of the Tuscan Archipelago. Results revealed and quantified the direct relationship between the temporary reduction of the maritime traffic due to the pandemic restrictions, and the probability of suffering damage from oil spill impact along the archipelago's rocky shores, which was highly context-dependent.

Spatial correlation reverses the compound effect of multiple stressors on rocky shore biofilm.

Understanding how multifactorial fluctuating environments affect species and communities remains one of the major challenges in ecology. The spatial configuration of the environment is known to generate complex patterns of correlation among multiple stressors. However, to what extent the spatial correlation between simultaneously fluctuating variables affects ecological assemblages in real-world conditions remains poorly understood. Here, we use field experiments and simulations to assess the influence of spatial correlation of two relevant climate variables - warming and sediment deposition following heavy precipitation - on the biomass and photosynthetic activity of rocky intertidal biofilm. First, we used a response-surface design experiment to establish the relation between biofilm, warming, and sediment deposition in the field. Second, we used the response surface to generate predictions of biofilm performance under different scenarios of warming and sediment correlation. Finally, we tested the predicted outcomes by manipulating the degree of correlation between the two climate variables in a second field experiment. Simulations stemming from the experimentally derived response surface showed how the degree and direction (positive or negative) of spatial correlation between warming and sediment deposition ultimately determined the nonlinear response of biofilm biomass (but not photosynthetic activity) to fluctuating levels of the two climate variables. Experimental results corroborated these predictions, probing the buffering effect of negative spatial correlation against extreme levels of warming and sediment deposition. Together, these results indicate that consideration of nonlinear response functions and local-scale patterns of correlation between climate drivers can improve our understanding and ability to predict ecological responses to multiple processes in heterogeneous environments.

The Obligate Symbiont "Candidatus Megaira polyxenophila" Has Variable Effects on the Growth of Different Host Species.

"Candidatus Megaira polyxenophila" is a recently described member of Rickettsiaceae which comprises exclusively obligate intracellular bacteria. Interestingly, these bacteria can be found in a huge diversity of eukaryotic hosts (protist, green algae, metazoa) living in marine, brackish or freshwater habitats. Screening of amplicon datasets revealed a high frequency of these bacteria especially in freshwater environments, most likely associated to eukaryotic hosts. The relationship of "Ca. Megaira polyxenophila" with their hosts and their impact on host fitness have not been studied so far. Even less is known regarding the responses of these intracellular bacteria to potential stressors. In this study, we used two phylogenetically close species of the freshwater ciliate Paramecium, Paramecium primaurelia and Paramecium pentaurelia (Ciliophora, Oligohymenophorea) naturally infected by "Ca. Megaira polyxenophila". In order to analyze the effect of the symbiont on the fitness of these two species, we compared the growth performance of both infected and aposymbiotic paramecia at different salinity levels in the range of freshwater and oligohaline brackish water i.e., at 0, 2, and 4.5 ppt. For the elimination of "Ca. Megaira polyxenophila" we established an antibiotic treatment to obtain symbiont-free lines and confirmed its success by fluorescence in situ hybridization (FISH). The population and infection dynamics during the growth experiment were observed by cell density counts and FISH. Paramecia fitness was compared applying generalized additive mixed models. Surprisingly, both infected Paramecium species showed higher densities under all salinity concentrations. The tested salinity concentrations did not significantly affect the growth of any of the two species directly, but we observed the loss of the endosymbiont after prolonged exposure to higher salinity levels. This experimental data might explain the higher frequency of "Ca. M. polyxenophila" in freshwater habitats as observed from amplicon data.

Direct observation of increasing recovery length before collapse of a marine benthic ecosystem.

Ecosystems can experience catastrophic transitions to alternative states, yet recent results have suggested that slowing down in rates of recovery after a perturbation may provide advance warning that a critical transition is approaching. Perturbation experiments with microbial populations have supported this hypothesis under controlled laboratory conditions, but evidence from natural ecosystems remains rare. Here, we manipulated rocky intertidal canopy algae to test the hypothesis that the spatial scale at which the system recovers from a perturbation in space should increase as the system approaches the tipping point, marking the transition from a canopy-dominated to a turf-dominated state. Empirical estimates of recovery length, a recently proposed spatial indicator of an approaching tipping point, were obtained by comparing the spatial scale at which algal turfs propagated into canopy-degraded regions with decreasing canopy cover. We show that recovery length increased along the gradient in canopy degradation, providing field-based evidence of spatial signatures of critical slowing down in natural conditions.