Forest mosaics, not savanna corridors, dominated in Southeast Asia during the Last Glacial Maximum.

The dominant paradigm is that large tracts of Southeast Asia's lowland rainforests were replaced with a "savanna corridor" during the cooler, more seasonal climates of the Last Glacial Maximum (LGM) (23,000 to 19,000 y ago). This interpretation has implications for understanding the resilience of Asia's tropical forests to projected climate change, implying a vulnerability to "savannization". A savanna corridor is also an important foundation for archaeological interpretations of how humans moved through and settled insular Southeast Asia and Australia. Yet an up-to-date, multiproxy, and empirical examination of the palaeoecological evidence for this corridor is lacking. We conducted qualitative and statistical analyses of 59 palaeoecological records across Southeast Asia to test the evidence for LGM savannization and clarify the relationships between methods, biogeography, and ecological change in the region from the start of Late Glacial Period (119,000 y ago) to the present. The pollen records typically show montane forest persistence during the LGM, while δ13C biomarker proxies indicate the expansion of C4-rich grasslands. We reconcile this discrepancy by hypothesizing the expansion of montane forest in the uplands and replacement of rainforest with seasonally dry tropical forest in the lowlands. We also find that smooth forest transitions between 34,000 and 2,000 y ago point to the capacity of Southeast Asia's ecosystems both to resist and recover from climate stressors, suggesting resilience to savannization. Finally, the timing of ecological change observed in our combined datasets indicates an 'early' onset of the LGM in Southeast Asia from ~30,000 y ago.

Marine reserves promote cycles in fish populations on ecological and evolutionary time scales.

Marine reserves are considered essential for sustainable fisheries, although their effectiveness compared to traditional fisheries management is debated. The effect of marine reserves is mostly studied on short ecological time scales, whereas fisheries-induced evolution is a well-established consequence of harvesting. Using a size-structured population model for an exploited fish population of which individuals spend their early life stages in a nursery habitat, we show that marine reserves will shift the mode of population regulation from low size-selective survival late in life to low, early-life survival due to strong resource competition. This shift promotes the occurrence of rapid ecological cycles driven by density-dependent recruitment as well as much slower evolutionary cycles driven by selection for the optimal body to leave the nursery grounds, especially with larger marine reserves. The evolutionary changes increase harvesting yields in terms of total biomass but cause disproportionately large decreases in yields of larger, adult fish. Our findings highlight the importance of carefully considering the size of marine reserves and the individual life history of fish when managing eco-evolutionary marine systems to ensure both population persistence as well as stable fisheries yields.

Regime shifts, trends, and variability of lake productivity at a global scale.

Lakes are often described as sentinels of global change. Phenomena like lake eutrophication, algal blooms, or reorganization in community composition belong to the most studied ecosystem regime shifts. However, although regime shifts have been well documented in several lakes, a global assessment of the prevalence of regime shifts is still missing, and, more in general, of the factors altering stability in lake status, is missing. Here, we provide a first global assessment of regime shifts and stability in the productivity of 1,015 lakes worldwide using trophic state index (TSI) time series derived from satellite imagery. We find that 12.8% of the lakes studied show regime shifts whose signatures are compatible with tipping points, while the number of detected regime shifts from low to high TSI has increased over time. Although our results suggest an overall stable picture for global lake dynamics, the limited instability signatures do not mean that lakes are insensitive to global change. Modeling the interaction between lake climatic, geophysical, and socioeconomic features and their stability properties, we find that the probability of a lake experiencing a tipping point increases with human population density in its catchment, while it decreases as the gross domestic product of that population increases. Our results show how quantifying lake productivity dynamics at a global scale highlights socioeconomic inequalities in conserving natural environments.

Consequences of kelp forest ecosystem shifts and predictors of persistence through multiple stressors.

Ecological communities can be stable over multiple generations, or rapidly shift into structurally and functionally different configurations. In kelp forest ecosystems, overgrazing by sea urchins can abruptly shift forests into alternative states that are void of macroalgae and primarily dominated by actively grazing sea urchins. Beginning in 2014, a sea urchin outbreak along the central coast of California resulted in a patchy mosaic of remnant forests interspersed with sea urchin barrens. In this study, we used a 14-year subtidal monitoring dataset of invertebrates, algae, and fishes to explore changes in community structure associated with the loss of forests. We found that the spatial mosaic of barrens and forests resulted in a region-wide shift in community structure. However, the magnitude of kelp forest loss and taxonomic-level consequences were spatially heterogeneous. Taxonomic diversity declined across the region, but there were no declines in richness for any group, suggesting compositional redistribution. Baseline ecological and environmental conditions, and sea urchin behaviour, explained the persistence of forests through multiple stressors. These results indicate that spatial heterogeneity in preexisting ecological and environmental conditions can explain patterns of community change.

Pyrogeography in flux: Reorganization of Australian fire regimes in a hotter world.

Changes to the spatiotemporal patterns of wildfire are having profound implications for ecosystems and society globally, but we have limited understanding of the extent to which fire regimes will reorganize in a warming world. While predicting regime shifts remains challenging because of complex climate-vegetation-fire feedbacks, understanding the climate niches of fire regimes provides a simple way to identify locations most at risk of regime change. Using globally available satellite datasets, we constructed 14 metrics describing the spatiotemporal dimensions of fire and then delineated Australia's pyroregions-the geographic area encapsulating a broad fire regime. Cluster analysis revealed 18 pyroregions, notably including the (1) high-intensity, infrequent fires of the temperate forests, (2) high-frequency, smaller fires of the tropical savanna, and (3) low-intensity, diurnal, human-engineered fires of the agricultural zones. To inform the risk of regime shifts, we identified locations where the climate under three CMIP6 scenarios is projected to shift (i) beyond each pyroregion's historical climate niche, and (ii) into climate space that is novel to the Australian continent. Under middle-of-the-road climate projections (SSP2-4.5), an average of 65% of the extent of the pyroregions occurred beyond their historical climate niches by 2081-2100. Further, 52% of pyroregion extents, on average, were projected to occur in climate space without present-day analogues on the Australian continent, implying high risk of shifting to states that also lack present-day counterparts. Pyroregions in tropical and hot-arid climates were most at risk of shifting into both locally and continentally novel climate space because (i) their niches are narrower than southern temperate pyroregions, and (ii) their already-hot climates lead to earlier departure from present-day climate space. Such a shift implies widespread risk of regime shifts and the emergence of no-analogue fire regimes. Our approach can be applied to other regions to assess vulnerability to rapid fire regime change.

Merging trait-based ecology and regime shift theory to anticipate community responses to warming.

Anthropogenic warming is altering species abundance, distribution, physiology, and more. How changes observed at the species level alter emergent community properties is an active and urgent area of research. Trait-based ecology and regime shift theory provide complementary ways to understand climate change impacts on communities, but these two bodies of work are only rarely integrated. Lack of integration handicaps our ability to understand community responses to warming, at a time when such understanding is critical. Therefore, we advocate for merging trait-based ecology with regime shift theory. We propose a general set of principles to guide this merger and apply these principles to research on marine communities in the rapidly warming North Atlantic. In our example, combining trait distribution and regime shift analyses at the community level yields greater insight than either alone. Looking forward, we identify a clear need for expanding quantitative approaches to collecting and merging trait-based and resilience metrics in order to advance our understanding of climate-driven community change.

Cooperation in public goods game does not require assortment and depends on population density.

The threshold public goods game is one of the best-known models of non-linear public goods dilemmas. Cooperators and defectors typically coexist in this game when the population is assumed to follow the so-called structured deme model. In this article, we develop a dynamical model of a general N-player game in which there is no deme structure: Individuals interact with randomly chosen neighbours and selection occurs between randomly chosen pairs of individuals. We show that in the deterministic limit, the dynamics in this model leads to the same replicator dynamics as in the structured deme model, i.e., coexistence of cooperators and defectors is typical in threshold public goods game even when the population is completely well mixed. We extend the model to study the effect of density dependence and density fluctuation on the dynamics. We show analytically and numerically that decreasing population density increases the equilibrium frequency of cooperators till the fixation of this strategy, but below a critical density cooperators abruptly disappear from the population. Our numerical investigations show that weak density fluctuations enhance cooperation, while strong fluctuations suppress it.

Fish microplastic ingestion may induce tipping points of aquatic ecosystems.

Microplastics can be ingested by a wide range of aquatic animals. Extensive studies have demonstrated that microplastic ingestion-albeit often not lethal-can affect a range of species life-history traits. However, it remains unclear how the sublethal effects of microplastics on individual levels scale up to influence ecosystem-level dynamics through cascading trophic interactions. Here we employ a well-studied, empirically fed three-species trophic chain model, which was parameterized to mimic a common type of aquatic ecosystems to examine how microplastic ingestion by fish on an intermediate trophic level can produce cascading effects on the species at both upper and lower trophic levels. We show that gradually increasing microplastics in the ingested substances of planktivorous fish may cause population structure effects such as skewed size distributions (i.e. reduced average body length vs. increased maximal body size), and induce abrupt declines in fish biomass and reproduction. Our model analysis demonstrates that these abrupt changes correspond to an ecosystem-level tipping point, crossing which difficult-to-reverse ecosystem degradation can happen. Importantly, microplastic pollution may interact with other anthropogenic stressors to reduce safe operating space of aquatic ecosystems. Our work contributes to better understanding complex effects of microplastic pollution and anticipating tipping points of aquatic ecosystems in a changing world. It also calls attention to an emerging threat that novel microplastic contaminants may lead to unexpected and abrupt degradation of aquatic ecosystems, and invites systematic studies on the ecosystem-level consequences of microplastic exposure.

First Passage Times of Long Transient Dynamics in Ecology.

Long transient dynamics in ecological models are characterized by extended periods in one state or regime before an eventual, and often abrupt, transition. One mechanism leading to long transient dynamics is the presence of ghost attractors, states where system dynamics slow down and the system lingers before eventually transitioning to the true attractor. This transition results solely from system dynamics rather than external factors. This paper investigates the dynamics of a classical herbivore-grazer model with the potential for ghost attractors or alternative stable states. We propose an intuitive threshold for first passage time analysis applicable to both bistable and ghost attractor regimes. By formulating the first passage time problem as a backward Kolmogorov equation, we examine how the mean first passage time changes as parameters are varied from the ghost attractor regime to the bistable one, through a saddle-node bifurcation. Our results reveal that the mean and variance of first passage times vary smoothly across the bifurcation threshold, eliminating the deterministic distinction between ghost attractors and bistable regimes. This work suggests that first passage time analysis can be an informative way to classify the length of a long transient. A better understanding of the duration of long transients may contribute to greater ecological understanding and more effective environmental management.

Effects of herbivorous fish on shallow lake ecosystems increase at moderate nutrient conditions.

Submerged macrophytes are vital in shallow lakes, as they provide critical ecosystem functions and services and can stabilize the clear-water conditions by various mechanisms. Nutrient enrichment reduces the resilience of macrophyte dominance in shallow lakes, thereby making them susceptible to shifts towards phytoplankton dominance following perturbations. Here, we conducted a mesocosm experiment to examine the individual and combined effects of nutrient enrichment and the addition of grass carp (Ctenopharyngodon idella) on the abundance of submerged macrophytes, epiphyton, and phytoplankton. We hypothesized that moderate nutrient enrichment facilitates macrophyte abundance, but also phytoplankton abundance after macrophyte removal by herbivorous fish. Our data showed that herbivory by grass carp could trigger a shift from macrophytes to algal dominance in mesocosms with moderate nutrient concentrations, but not in those with low nutrient concentrations. Moderate nutrient enrichment alone promoted submerged macrophyte growth, whereas the introduction of grass carp induced a collapse of submerged macrophytes regardless of nutrient conditions. Moreover, the introduction of grass carp showed more negative effects on light conditions of the water column in mesocosms with moderate nutrient concentrations compared to those with low nutrient concentrations. A recovery of submerged macrophytes might thus be limited by low light availability in lakes with moderate nutrient conditions suffering grass carp perturbation. Our results suggest that submerged macrophyte-dominated shallow lakes with moderate nutrient conditions are vulnerable to perturbation by herbivorous fish such as grass carp. In turn, managing the abundance of herbivores in these lakes can support the dominance of macrophytes and associated clear water conditions.

Functional diversity can facilitate the collapse of an undesirable ecosystem state.

Biodiversity may increase ecosystem resilience. However, we have limited understanding if this holds true for ecosystems that respond to gradual environmental change with abrupt shifts to an alternative state. We used a mathematical model of anoxic-oxic regime shifts and explored how trait diversity in three groups of bacteria influences resilience. We found that trait diversity did not always increase resilience: greater diversity in two of the groups increased but in one group decreased resilience of their preferred ecosystem state. We also found that simultaneous trait diversity in multiple groups often led to reduced or erased diversity effects. Overall, our results suggest that higher diversity can increase resilience but can also promote collapse when diversity occurs in a functional group that negatively influences the state it occurs in. We propose this mechanism as a potential management approach to facilitate the recovery of a desired ecosystem state.

Eutrophication decreases ecological resilience by reducing species diversity and altering functional traits of submerged macrophytes.

Positive feedback is key to producing alternative stable states and largely determines ecological resilience in response to external perturbations. Understanding the positive feedback mechanisms in macrophyte-dominated lakes is crucial for resilience-based management and restoration. Based on the field investigation of submerged macrophyte communities in 35 lakes in China, we found that morphological complexity (MC) and morphological plasticity (MP) are correlated with the stoichiometric homeostasis of phosphorus (HP ) and are related to ecosystem structure, functioning, and stability. We also found that the positive feedback strength of lakes dominated by macrophytes is biomass- and diversity-dependent. Eutrophication can decrease the community biomass by decreasing community MC, MP, and HP and the species diversity through low-light availability, ultimately decreasing the positive feedback strength and resilience of clear water states. We argue that functional traits and species diversity should be considered to build more resilient ecosystems in future changing environment scenarios.

Effects of interactions between anthropogenic stressors and recurring perturbations on ecosystem resilience and collapse.

Insights into declines in ecosystem resilience and their causes and effects can inform preemptive action to avoid ecosystem collapse and loss of biodiversity, ecosystem services, and human well-being. Empirical studies of ecosystem collapse are rare and hampered by ecosystem complexity, nonlinear and lagged responses, and interactions across scales. We investigated how an anthropogenic stressor could diminish ecosystem resilience to a recurring perturbation by altering a critical ecosystem driver. We studied groundwater-dependent, peat-accumulating, fire-prone wetlands known as upland swamps in southeastern Australia. We hypothesized that underground mining (stressor) reduces resilience of these wetlands to landscape fires (perturbation) by diminishing groundwater, a key ecosystem driver. We monitored soil moisture as an indicator of ecosystem resilience during and after underground mining. After landscape fire, we compared responses of multiple state variables representing ecosystem structure, composition, and function in swamps within the mining footprint with unmined reference swamps. Soil moisture declined without recovery in swamps with mine subsidence (i.e., undermined), but was maintained in reference swamps over 8 years (effect size 1.8). Relative to burned reference swamps, burned undermined swamps showed greater loss of peat via substrate combustion; reduced cover, height, and biomass of regenerating vegetation; reduced postfire plant species richness and abundance; altered plant species composition; increased mortality rates of woody plants; reduced postfire seedling recruitment; and extirpation of a hydrophilic animal. Undermined swamps therefore showed strong symptoms of postfire ecosystem collapse, whereas reference swamps regenerated vigorously. We found that an anthropogenic stressor diminished the resilience of an ecosystem to recurring perturbations, predisposing it to collapse. Avoidance of ecosystem collapse hinges on early diagnosis of mechanisms and preventative risk reduction. It may be possible to delay or ameliorate symptoms of collapse or to restore resilience, but the latter appears unlikely in our study system due to fundamental alteration of a critical ecosystem driver. Efectos de las interacciones entre los estresantes antropogénicos y las perturbaciones recurrentes sobre la resiliencia y el colapso de los ecosistemas.

La comprensión de la declinación en la resiliencia de los ecosistemas y sus causas y efectos puede orientar las acciones preventivas para evitar el colapso ecosistémico y la pérdida de biodiversidad, servicios ambientales y bienestar humano. Los estudios empíricos del colapso ecosistémico son escasos y se enfrentan a obstáculos como la complejidad del ecosistema, respuestas rezagadas y no lineales e interacciones entre las escalas. Investigamos cómo un estresante antropogénico podría reducir la resiliencia del ecosistema a una perturbación recurrente mediante la alteración de un causante importante. Estudiamos los humedales dependientes de aguas subterráneas que acumulan turbas y son propicios a incendios conocidos como pantanos de tierras altas en el sureste de Australia. Nuestra hipótesis fue que la minería subterránea (estresante) reduce la resiliencia de estos humedales a incendios (perturbación) al disminuir el agua subterránea, un causante clave para el ecosistema. Monitoreamos la humedad del suelo como un indicador de la resiliencia del ecosistema durante y después de la minería subterránea. Después de los incendios, comparamos la respuesta de múltiples variables de estado que representaban la estructura, composición y función del ecosistema en los pantanos dentro de la huella minera con los pantanos referenciales sin minería. La humedad del suelo declinó sin recuperación en los pantanos con hundimientos mineros (es decir, socavones) pero se mantuvo en los pantanos referenciales durante ocho años (tamaño del efecto: 1.8). En relación a los pantanos referenciales incendiados, los pantanos con socavones e incendios mostraron una mayor pérdida de turba mediante la combustión del sustrato; reducción en la cobertura, altura y regeneración de biomasa de la vegetación; reducción en la riqueza y abundancia de especies vegetales post incendio; alteraciones en la composición de especies vegetales; incremento en la mortalidad de las plantas leñosas; reducción en el reclutamiento post incendio de plántulas; y la extirpación de un animal hidrofílico. Por lo tanto, los pantanos con socavones mostraron síntomas fuertes de un colapso ecosistémico post incendio, mientras que los pantanos referenciales se regeneraron vigorosamente. Descubrimos que los estresantes antropogénicos redujeron la resiliencia de un ecosistema a perturbaciones recurrentes, lo que lo predispone al colapso. La eliminación de este colapso depende de un diagnóstico temprano de mecanismos y reducción del riesgo preventivo. Puede ser posible retardar o mitigar los síntomas del colapso o restaurar la resiliencia, aunque lo último parece ser improbable en nuestro sistema de estudio debido a la alteración fundamental de un causante importante del ecosistema.

The emergence and resilience of self-organized governance in coupled infrastructure systems.

Studies of small-scale, self-organized social-ecological systems have contributed to our understanding of successful governance of shared resources. However, the lack of formal analytically tractable models of such coupled infrastructure systems makes it difficult to connect this understanding to such concepts as stability, robustness, and resilience, which are increasingly important in considering such systems. In this paper, we mathematically operationalize a widely used conceptual framework via a stylized dynamical model. The model yields a wide range of system outcomes: sustainability or collapse, infrastructure at full or partial capacity, and social agents seeking outside opportunities or exclusively engaging in the system. The low dimensionality of the model enables us to derive these conditions in clear relationships of biophysical and social factors describing the coupled system. Analysis of the model further reveals regime shifts, trade-offs, and potential pitfalls that one may face in governing these self-organized systems. The intuition and insights derived from the model lay ground for more rigorous treatment of robustness and resilience of self-organized coupled infrastructure systems, which can lead to more effective governance.

Disentangling the effects of phosphorus loading on food web stability in a large shallow lake.

Excessive nutrient loads reduce ecosystem resilience, resulting in fundamental changes in ecosystem structure and function when exceeding a certain threshold. However, quantitative analysis of the processes by which nutrient loading affects ecosystem resilience requires further exploration. Food web stability is at the heart of ecosystem resilience. In this study, we simulated the dynamics of the food web under different phosphorus loads for Lake Baiyangdian using the PCLake model and calculated the food web stability. Our results showed that there was a good correspondence between the food web stability and ecosystem state response to phosphorus loads. This relationship confirmed that food web stability could be regarded as a signal for the state transition in a real lake ecosystem. Moreover, our estimates suggested that food web stability was influenced only by several functional groups and their interaction strength. Diatoms and zooplankton were the key functional groups that affected food web stability. Phosphorus loads alter the distribution of functional group biomass, which in turn affects energy delivery and, ultimately, the stability of the food web. Corresponding to functional groups, the interactions among zooplankton, diatoms and detritus had the greatest impact, and the interaction strength of the three was positively correlated with food web stability. Overall, our study explained that food-web stability was critical to characterize ecosystem resilience response to external disturbances and can be turned into a scientific tool for lake ecosystem management.

Reassessment of the declines in the largest freshwater lake in China (Poyang Lake): uneven trends, risks and underlying causes.

Water level decline in the largest freshwater lake in China (Poyang Lake) has raised grave concerns and continuous debates regarding the proposal of a water control project. Previous hydrologic investigations on the water level decline in Poyang Lake were conducted mostly during recession season and typical dry years and hence lacked the comprehensive view of associated risks and possible spatial heterogeneity of trend during low water level periods. The present study reassessed the long-term trend and regime shift of low water level variations and their associated risks based on hydrological data across multiple stations in Poyang Lake during 1952-2021. The underlying causes of the trends of water level decline were further investigated. Results showed uneven trends and risks of water level variations in different seasons and across different lake regions. The water level of all five hydrological stations in Poyang Lake significantly dropped in the recession season, and the risks of water level decline evidently increased since 2003, which could be largely attributed to the water level drop in the Yangtze River. In terms of the dry season, clear spatial differences of long-term trend of water level were found, with water level in the central and southern lake regions significantly dropping, which was probably caused by dramatic bathymetry undercutting in the central and northern lake regions. In addition, the impacts of topographic changes became significant when water level of Hukou fell below 13.8 and 11.8m for the northern and southern lake regions, respectively. By contrast, water level in the northern lake region showed increasing trends in the dry season. In addition, only the occurrence time of water level under moderate risk significantly advanced for all stations except for Hukou. The present study could provide a full picture of low water level trends, associated risks across different lake regions and underlying causes in Poyang Lake, thus providing insights into adaptive water resources management.

Evaluating the performance of temporal and spatial early warning statistics of algal blooms.

Regime shifts have large consequences for ecosystems and the services they provide. However, understanding the potential for, causes of, proximity to, and thresholds for regime shifts in nearly all settings is difficult. Generic statistical indicators of resilience have been proposed and studied in a wide range of ecosystems as a method to detect when regime shifts are becoming more likely without direct knowledge of underlying system dynamics or thresholds. These early warning statistics (EWS) have been studied separately but there have been few examples that directly compare temporal and spatial EWS in ecosystem-scale empirical data. To test these methods, we collected high-frequency time series and high-resolution spatial data during a whole-lake fertilization experiment while also monitoring an adjacent reference lake. We calculated two common EWS, standard deviation and autocorrelation, in both time series and spatial data to evaluate their performance prior to the resulting algal bloom. We also applied the quickest detection method to generate binary alarms of resilience change from temporal EWS. One temporal EWS, rolling window standard deviation, provided advanced warning in most variables prior to the bloom, showing trends and between-lake patterns consistent with theory. In contrast, temporal autocorrelation and both measures of spatial EWS (spatial SD, Moran's I) provided little or no warning. By compiling time series data from this and past experiments with and without nutrient additions, we were able to evaluate temporal EWS performance for both constant and changing resilience conditions. True positive alarm rates were 2.5-8.3 times higher for rolling window standard deviation when a lake was being pushed towards a bloom than the rate of false positives when it was not. For rolling window autocorrelation, alarm rates were much lower and no variable had a higher true positive than false positive alarm rate. Our findings suggest temporal EWS provide advanced warning of algal blooms and that this approach could help managers prepare for and/or minimize negative bloom impacts.

Tracking spatial regimes as an early warning for a species of conservation concern.

In this era of global environmental change and rapid regime shifts, managing core areas that species require to survive and persist is a grand challenge for conservation. Wildlife monitoring data are often limited or local in scale. The emerging ability to map and track spatial regimes (i.e., the spatial manifestation of state transitions) using advanced geospatial vegetation data has the potential to provide earlier warnings of habitat loss because many species of conservation concern strongly avoid spatial regime boundaries. Using 23 yr of data for the lek locations of Greater Prairie-Chicken (Tympanuchus cupido; GPC) in a remnant grassland ecosystem, we demonstrate how mapping changes in the boundaries between grassland and woodland spatial regimes provide a spatially explicit early warning signal for habitat loss for an iconic and vulnerable grassland-obligate known to be highly sensitive to woody plant encroachment. We tested whether a newly proposed metric for the quantification of spatial regimes captured well-known responses of GPC to woody plant expansion into grasslands. Resource selection functions showed that the grass:woody spatial regime boundary strength explained the probability of 80% of relative lek occurrence, and GPC strongly avoided grass:woody spatial regime boundaries at broad scales. Both findings are consistent with well-known expectations derived from GPC ecology. These results provide strong evidence for vegetation-derived delineations of spatial regimes to serve as generalized signals of early warning for state transitions that have major consequences to biodiversity conservation. Mapping spatial regime boundaries over time provided interpretable early warnings of habitat loss. Woody plant regimes displaced grassland regimes starting from the edges of the study area and constricting inward. Correspondingly, the relative probability of lek occurrence constricted in space. Similarly, the temporal trajectory of spatial regime boundary strength increased over time and moved closer to the observed limit of GPC lek site usage relative to grass:woody boundary strength. These novel spatial metrics allow managers to rapidly screen for early warning signals of spatial regime shifts and adapt management practices to defend and grow habitat cores at broad scales.

Analysis of the onset of a regime shift and detecting early warning signs of major population changes in a two-trophic three-species predator-prey model with long-term transients.

Identifying early warning signs of sudden population changes and mechanisms leading to regime shifts are highly desirable in population biology. In this paper, a two-trophic ecosystem comprising of two species of predators, competing for their common prey, with explicit interference competition is considered. With proper rescaling, the model is portrayed as a singularly perturbed system with fast prey dynamics and slow dynamics of the predators. In a parameter regime near singular Hopf bifurcation, chaotic mixed-mode oscillations (MMOs), featuring concatenation of small and large amplitude oscillations are observed as long-lasting transients before the system approaches its asymptotic state. To analyze the dynamical cause that initiates a large amplitude oscillation in an MMO orbit, the model is reduced to a suitable normal form near the singular-Hopf point. The normal form possesses a separatrix surface that separates two different types of oscillations. A large amplitude oscillation is initiated if a trajectory moves from the "inner" to the "outer side" of this surface. A set of conditions on the normal form variables are obtained to determine whether a trajectory would exhibit another cycle of MMO dynamics before experiencing a regime shift (i.e. approaching its asymptotic state). These conditions serve as early warning signs for a sudden population shift as well as detect the onset of a regime shift in this ecological model.

Climate-driven regime shifts in a mangrove-salt marsh ecotone over the past 250 years.

Climate change is driving the tropicalization of temperate ecosystems by shifting the range edges of numerous species poleward. Over the past few decades, mangroves have rapidly displaced salt marshes near multiple poleward mangrove range limits, including in northeast Florida. It is uncertain whether such mangrove expansions are due to anthropogenic climate change or natural climate variability. We combined historical accounts from books, personal journals, scientific articles, logbooks, photographs, and maps with climate data to show that the current ecotone between mangroves and salt marshes in northeast Florida has shifted between mangrove and salt marsh dominance at least 6 times between the late 1700s and 2017 due to decadal-scale fluctuations in the frequency and intensity of extreme cold events. Model projections of daily minimum temperature from 2000 through 2100 indicate an increase in annual minimum temperature by 0.5 °C/decade. Thus, although recent mangrove range expansion should indeed be placed into a broader historical context of an oscillating system, climate projections suggest that the recent trend may represent a more permanent regime shift due to the effects of climate change.