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The temperature-size rule is one of the universal rules in ecology and states that ectotherms in warmer waters will grow faster as juveniles, mature at smaller sizes and younger ages, and reach smaller maximum body sizes. Many models have unsuccessfully attempted to reproduce temperature-size rule-consistent life histories by using two-term (anabolism and catabolism) Pütter-type growth models, such as the von Bertalanffy. Here, we present a physiologically structured individual growth model, which incorporates an energy budget and optimizes energy allocation to growth, reproduction, and reserves. Growth, maturation, and reproductive output emerge as a result of life-history optimization to specific physiological rates and mortality conditions. To assess which processes can lead to temperature-size rule-type life histories, we simulate 42 scenarios that differ in temperature and body size dependencies of intake, metabolism, and mortality rates. Results show that the temperature-size rule can emerge in two ways. The first way requires both intake and metabolism to increase with temperature, but the temperature-body size interaction of the two rates must lead to relatively faster intake increase in small individuals and relatively larger metabolism increase in large ones. The second way requires only higher temperature-driven natural mortality and faster intake rates in early life (no change in metabolic rates is needed). This selects for faster life histories with earlier maturation and increased reproductive output. Our model provides a novel mechanistic and evolutionary framework for identifying the conditions necessary for the temperature-size rule. It shows that the temperature-size rule is likely to reflect both physiological changes and life-history optimization and that use of von Bertalanffy-type models, which do not include reproduction processes, can hinder our ability to understand and predict ectotherm responses to climate change.

Fish and other ectotherms living in warmer waters often grow faster as juveniles, mature earlier, but become smaller adults. Known as the temperature-size rule (TSR), this pattern is commonly attributed to higher metabolism in warmer waters, leaving fewer resources for growth. An alternative explanation focuses on growth and reproduction trade-offs across temperatures. We tested these hypotheses by measuring growth, maturation, metabolism and reproductive allocation from zebrafish populations kept at 26 and 30°C across six generations. Zebrafish growth and maturation followed TSR expectations but were not explained by baseline metabolic rate, which converged between temperature treatments after a few generations. Rather, we found that females at 30°C allocated more to reproduction, especially when maturing at the smallest sizes. We show that elevated temperatures do not necessarily increase baseline metabolism if sufficient acclimation is allowed and call for an urgent revision of modelling assumptions used to predict population and ecosystem responses to warming.

The northern pike (Esox lucius) is an iconic predatory fish species of significant recreational value and ecological role in the Baltic Sea. Some earlier studies indicate local declines of pike in the region, but a thorough spatial evaluation of regional population trends of pike in the Baltic Sea is lacking. In this study, we collate data from 59 unique time-series from fisheries landings and fishery-independent monitoring programs to address temporal trends in pike populations since the mid-2000′s in eight countries surrounding the Baltic Sea. In a common analysis considering all time-series in concert, we found indications of an overall regional temporal decline of pike in the Baltic Sea, but trends differed among countries. Individual negative trends in time-series were moreover found in several regions of the Baltic Sea, but predominantly so in the central and southern parts, while positive trends were only found in Estonia and northern Finland. The mix of data used in this study is inherently noisy and to some extent of uncertain quality, but as a result of the overall negative trends, together with the socioeconomic and ecological importance of pike in coastal areas of the Baltic Sea, we suggest that actions should be taken to protect and restore pike populations. Management measures should be performed in combination with improved fishery-independent monitoring programs to provide data of better quality and development of citizen-science approaches as a data source for population estimates. Possible measures that could strengthen pike populations include harvest regulations (including size limits, no-take areas and spawning closures), habitat protection and restoration, and an ecosystem-based approach to management considering also the impact of natural predators.

Most fish populations around the world are unassessed and their status is unknown. Size based methods could provide fast and transparent assessments, but they require information on fish sizes. Citizen science programs, social media and smart phone applications generate millions of georeferenced fish images globally. Machine learning based fish species and size identification could help turn these images into valuable data for population status assessments. We present a machine learning, image classification based method to identify fish size classes from photos of anglers holding fish. To train the model we group images into ten 5-10 cm size classes, similar to classes used in underwater visual fish surveys. The model was trained using 2602 images from angler citizen science platforms MyCatch and FishSizeProject. Although the number of images was limited, the model achieved an overall accuracy of ~50%. Importantly, the misidentification of size classes was consistent across 20 separate model training rounds, each conducted with an independent, random allocation of images for training and test datasets. Our method suggests that photo based fish size class identification is feasible, and that prediction uncertainty should be incorporated into subsequent analysis, as it is done with fish ageing errors in fisheries stock assessments.

Pikeperch (Sander lucioperca) is a European fresh and brackish water piscivorous fish, important as both a key predator and a valuable commercial and recreational fisheries species. There are concerns that some stocks are depleted due to overfishing and environmental changes. We review data collection and population assessments currently used for nine pikeperch stocks across six European countries and apply a unified assessment framework to evaluate population status and trends. For this we first standardised commercial, scientific, and recreational catch-per-unit-effort (CPUE) and catch time series and then applied Bayesian surplus production models. Our results showed that three stocks (including two in the Baltic Sea) were strongly depleted, with estimated biomasses considerably lower than the biomass at maximum sustainable yield (Bmsy). Other stocks were either close or higher than their estimated Bmsy. Looking at the trends, we find that four stocks (Lake Oulujärvi, Kvädöfjärden, Lake Peipsi and Lipno) showed increasing biomass trends and two (Curonian Lagoon, Galtfjärden) had a strong decline in biomass. In most cases the stocks with clear signs of recovery were also those for which strong management strategies have been implemented. We find that, despite pikeperch being one of the most valuable inland fisheries, formalised stock assessments and regular surveys remain rare. Importantly, although most stocks are strongly targeted by recreational fishing, estimates of recreational catch are highly uncertain. We conclude that data limited stock assessment methods are useful for assessing fish population status and highlight an urgent need to improve pikeperch scientific monitoring and assessment of recreational catches.

It is manifest that COVID-19 lockdowns extensively impacted human interactions with natural ecosystems. One example is recreational fishing, an activity which involves nearly 1 in 10 people in developed countries. Fishing licence sales and direct observations at popular angling locations suggest that recreational fishing effort increased substantially during lockdowns. However, the extent and duration of this increase remain largely unknown due to a lack of objective data. We used four years (2018 to 2021) of anonymous, high-resolution data from a personal fish-finder device to explore the impact of COVID-19 lockdowns on recreational fishing effort in four European countries (Lithuania, the Czech Republic, Denmark, and Germany). We show that device use and, by extension, angling effort increased 1.2-3.8 fold during March-May 2020 and remained elevated even at the end of 2021 in all countries except Denmark. Fishing during the first lockdown also became more frequent during weekdays. Statistical models with the full set of fixed (weekdays, lockdown, population) and random (season, year, administrative unit) factors typically explained 50-70% of the variation, suggesting that device use and angling effort were relatively consistent and predictable through space and time. Our study demonstrates that recreational fishing behaviour can change substantially and rapidly in response to societal shifts, with profound ecological, human well-being and economic implications. We also show the potential of angler devices and smartphone applications to supply data for high-resolution fishing effort analysis and encourage more extensive science and industry collaborations to take advantage of this information.Significance statementRecreational fishing is a popular and widespread activity with ecological, social and economic impacts, though problematic to assess and manage due to a paucity of information regarding effort and catch. Here, we use high-resolution data from a personal angler sonar device to show how the COVID-19 pandemic changed angler behaviour and fishing effort across Europe. We demonstrate that angling effort doubled and remained higher at the end of 2021 than before the first lockdowns. Such rapid and profound changes could have significant consequences for aquatic ecosystems, possibly requiring new management approaches. We encourage the adoption of novel data from angler devices, citizen science, and more active science-industry collaborations to improve recreational fishing assessment and management.

Glacial relict mysid and amphipod crustaceans are characterised by their affinity for cold and well-oxygenated waters and inability to disperse upstream or by external agents. These crustaceans occur in large and deep lakes of Northern and Central Europe and North America, their distributions shaped by glaciation events. In Europe, along the southern edge of their distribution (Germany, Poland, Lithuania, Belarus), glacial relict crustaceans are threatened by eutrophication and global warming.This study assesses the status of three glacial relict malacostracan species in Lithuania; the amphipodsMonoporeia affinisandPallaseopsis quadrispinosa, and mysidMysis relicta, and models their abundance as a function of environmental variables and the presence of invasive Ponto-Caspian mysids and amphipods.Our results revealed thatM. affinisis likely extinct in the country, whereasM. relictawas found in only 9 out of 16 locations from which it was previously recorded. The distribution ofP. quadrispinosaappears to be shrinking.Lake depth and water flowthrough intensity were significantly and positively associated with the relative abundance of relict mysids and amphipods, but no association was found with lake size or the presence of invasive Ponto-Caspian crustaceans.We conclude that urgent action to mitigate the effects of nutrient run-off is needed to improve the status of glacial relict and other species that require good water quality. We also propose the re-introduction of glacial relict species in Lake Drūkšiai, where they went extinct during the operation of the Ignalina nuclear power plant that heated the lake, but where deep-water environmental conditions have improved following the powerplant closure in 2010.

Citizen science platforms, social media and smart phone applications enable the collection of large amounts of georeferenced images. This provides a huge opportunity in biodiversity and ecological research, but also creates challenges for efficient data handling and processing. Recreational and small-scale fisheries is one of the fields that could be revolutionised by efficient, widely accessible and machine learning-based processing of georeferenced images. Most non-commercial inland and coastal fisheries are considered data poor and are rarely assessed, yet they provide multiple societal benefits and can have substantial ecological impacts. Given that large quantities of georeferenced fish images are being collected by fishers every day, artificial intelligence (AI) and computer vision applications offer a great opportunity to automate their analyses by providing species identification, and potentially also fish size estimation. This would deliver data needed for fisheries management and fisher engagement. To date, however, many AI image analysis applications in fisheries are focused on the commercial sector, limited to specific species or settings, and are not publicly available. In addition, using AI and computer vision tools often requires a strong background in programming. In this study, we aim to facilitate broader use of computer vision tools in fisheries and ecological research by compiling an open-source user friendly and modular framework for large-scale image storage, handling, annotation and automatic classification, using cost- and labour-efficient methodologies. The tool is based on TensorFlow Lite Model Maker library, and includes data augmentation and transfer learning techniques applied to different convolutional neural network models. We demonstrate the potential application of this framework using a small example dataset of fish images taken through a recreational fishing smartphone application. The framework presented here can be used to develop region-specific species identification models, which could potentially be combined into a larger hierarchical model.

Oxygen bioavailability is declining in aquatic systems worldwide as a result of climate change and other anthropogenic stressors. For aquatic organisms, the consequences are poorly known but are likely to reflect both direct effects of declining oxygen bioavailability and interactions between oxygen and other stressors, including two—warming and acidification— that have received substantial attention in recent decades and that typically accompany oxygen changes. Drawing on the collected papers in this symposium volume (“An Oxygen Perspective on Climate Change”), we outline the causes and consequences of declining oxygen bioavailability. First, we discuss the scope of natural and predicted anthropogenic changes in aquatic oxygen levels. Although modern organisms are the result of long evolutionary histories during which they were exposed to natural oxygen regimes, anthropogenic change is now exposing them to more extreme conditions and novel combinations of low oxygen with other stressors. Second, we identify behavioral and physiological mechanisms that underlie the interactive effects of oxygen with other stressors, and we assess the range of potential organismal responses to oxygen limitation that occur across levels of biological organization and over multiple timescales. We argue that metabolism and energetics provide a powerful and unifying framework for understanding organism-oxygen interactions. Third, we conclude by out-lining a set of approaches for maximizing the effectiveness of future work, including focusing on long-term experiments using biologically realistic variation in experimental factors and taking truly cross-disciplinary and integrative approaches to understanding and predicting future effects.

Citizen science platforms, social media and multiple smart phone applications enable collection of large amounts of georeferenced images. This provides a huge opportunity in biodiversity and ecological research, but also creates challenges for efficient data handling and processing. Recreational and small-scale fisheries is one of the fields that could be revolutionised by efficient, widely accessible and machine learning based processing of georeferenced images. The majority of non-commercial inland and coastal fisheries are considered data poor and are rarely assessed, yet they provide multiple societal benefits and can have large ecological impacts. Given that large quantities of fish observations and images are being collected by fishers every day, artificial intelligence (AI) and computer vision applications offer a great opportunity to improve data collection, automate analyses and inform management. Yet, to date, many AI image analysis applications in fisheries are focused on the commercial sector and are not publicly available for community use. In this study we present an open-source modular framework for large scale image storage, handling, annotation and automatic classification, using cost- and labour-efficient methodologies. The tool is based on TensorFlow Lite Model Maker library and includes data augmentation and transfer learning techniques, applied to different convolutional neural network models. We demonstrate the implementation of this framework in an example case study for automatic fish species identification from images taken through a recreational fishing smartphone application. The framework presented here is highly customisable for further advancement and community based image collection and annotation.

Changing sea temperatures and primary productivity are rapidly altering marine ecosystems, but with considerable uncertainty in our understanding of the relative importance of these drivers and how their interactions may affect fisheries yield through complex food webs. Such outcomes are more difficult to predict for shallow coastal ecosystems than those in pelagic and shelf habitats, because coastal food webs are fuelled by a combination of separate pelagic and benthic energy pathways. Using long-term, empirical field data, we developed a novel multispecies size spectrum model for shallow coastal reefs. We include size-structured benthic and pelagic resources and trophic structures, allowing us to explore potential climate change scenarios that involve varying combinations of warming with changes in benthic and pelagic resources. Our model predicts that changes in resource levels will have much stronger impacts on fish biomass and yields than changes driven by physiological responses to temperature. Under increased plankton abundance, species in all trophic groups were predicted to increase in biomass, average size and yields. By contrast, changes in benthic resource produced variable responses across coastal trophic groups. Increased benthic resource led to increasing benthivorous and piscivorous fish biomass, yields and mean body sizes, but decreases in herbivores and planktivores. When resource changes were combined with warming seas, physiological responses generally decreased species’ biomass and yields. Our results suggest that the source, size and abundance of primary and secondary producers are critical to understanding impacts of warming seas on coastal fish communities. Understanding changes in benthic production and its implications for coastal fisheries requires urgent attention. Our modified size spectrum model provides a framework for further study of benthic and pelagic energy pathways that can be easily adapted to other ecosystems.

Fishing is a strong selective force and is supposed to select for earlier maturation at smaller body size. However, the extent to which fishing‐induced evolution is shaping ecosystems remains debated. This is in part because it is challenging to disentangle fishing from other selective forces (e.g., size‐structured predation and cannibalism) in complex ecosystems undergoing rapid change.Changes in maturation size from fishing and predation have previously been explored with multi‐species physiologically structured models but assumed separation of ecological and evolutionary timescales. To assess the eco‐evolutionary impact of fishing and predation at the same timescale, we developed a stochastic physiologically size‐structured food‐web model, where new phenotypes are introduced randomly through time enabling dynamic simulation of species' relative maturation sizes under different types of selection pressures.Using the model, we carried out a fully factorial in silico experiment to assess how maturation size would change in the absence and presence of both fishing and predation (including cannibalism). We carried out ten replicate stochastic simulations exposed to all combinations of fishing and predation in a model community of nine interacting fish species ranging in their maximum sizes from 10 g to 100 kg. We visualized and statistically analyzed the results using linear models.The effects of fishing on maturation size depended on whether or not predation was enabled and differed substantially across species. Fishing consistently reduced the maturation sizes of two largest species whether or not predation was enabled and this decrease was seen even at low fishing intensities (F = 0.2 per year). In contrast, the maturation sizes of the three smallest species evolved to become smaller through time but this happened regardless of the levels of predation or fishing. For the four medium‐size species, the effect of fishing was highly variable with more species showing significant and larger fishing effects in the presence of predation.Ultimately our results suggest that the interactive effects of predation and fishing can have marked effects on species' maturation sizes, but that, at least for the largest species, predation does not counterbalance the evolutionary effect of fishing. Our model also produced relative maturation sizes that are broadly consistent with empirical estimates for many fish species., Fishing is a strong evolutionary force, but its interplay with predation remains unclear. Using a community size‐spectrum model we show that fishing has strongest evolutionary effect on large predatory species, but minor impact on small species. Middle‐sized species are sandwiched between fishing and predation and their responses are highly variable.

The size and age at which individuals mature is rapidly changing due to plastic and evolved responses to fisheries harvest and global warming. Understanding the nature of these changes is essential because maturity schedules are critical in determining population demography and ultimately, the economic value and viability of fisheries. Detecting maturity changes is, however, practically difficult and costly. A recently proposed biphasic growth modelling likelihood profiling method offers great potential as it can statistically estimate age-at-maturity from population-level size-at-age data, using the change-point in growth that occurs at maturity. Yet, the performance of the method on typical marine fisheries datasets remains untested. Here, we assessed the suitability of 12 North Sea and Australian species’ datasets for the likelihood profiling approach. The majority of the fisheries datasets were unsuitable as they had too small sample sizes or too large size-at-age variation. Further, datasets that did satisfy data requirements generally showed no correlation between empirical and model-derived maturity estimates. To understand why the biphasic approach had low performance we explored its sensitivity using simulated datasets. We found that method performance for marine fisheries datasets is likely to be low because of: (1) truncated age structures due to intensive fishing, (2) an under-representation of young individuals in datasets due to common fisheries-sampling protocols, and (3) large intrapopulation variability in growth curves. To improve our ability to detect maturation changes from population level size-at-age data we need to improve data collection protocols for fisheries monitoring.

Climate‐driven species redistribution is pervasive and accelerating, yet the complex mechanisms at play remain poorly understood. The implications of large‐scale species redistribution for natural systems and human societies have resulted in a large number of studies exploring the effects on individual species and ecological communities worldwide. Whilst many studies have investigated discrete components of species redistribution, the integration required for a more complete mechanistic understanding is lacking. In this paper, we provide a framework for synthesising approaches to more robustly understand and predict marine species redistributions. We conceptualise the stages and processes involved in climate‐driven species redistribution at increasing levels of biological organisation, and synthesize the laboratory, field and modelling approaches used to study redistribution related processes at individual, population and community levels. We then summarise links between scales of biological organisation and methodological approaches in a hierarchical framework that represents an integrated mechanistic assessment of climate‐driven species redistributions. In a rapidly expanding field of research, this framework provides direction for: 1) guiding future research, 2) highlighting key knowledge gaps, 3) fostering data exchange and collaboration between disciplines and 4) improving shared capacity to predict and therefore, inform the proactive management of climate impacts on natural systems.

Ectotherms generally shrink under experimental warming, but whether this pattern extends to wild populations is uncertain. We analysed ten million visual survey records, spanning the Australian continent and multiple decades and comprising the most common coastal reef fishes (335 species). We found that temperature indeed drives spatial and temporal changes in fish body size, but not consistently in the negative fashion expected. Around 55% of species were smaller in warmer waters (especially among small-bodied species), while 45% were bigger. The direction of a species’ response to temperature through space was generally consistent with its response to temperature increase through time at any given location, suggesting that spatial trends could help forecast fish responses to long-term warming. However, temporal changes were about ten times faster than spatial trends (~4% versus ~40% body size change per 1 °C change through space and time, respectively). The rapid and variable responses of fish size to warming may herald unexpected impacts on ecosystem restructuring, with potentially greater consequences than if all species were shrinking.

The Ponto-Caspian region is an endemicity hotspot that harbors several crustacean radiations, among which amphipods are the most diverse. These poorly known species are severely threatened in their native range, while at the same time they are invading European inland waters with significant ecological consequences. A proper taxonomic knowledge of this fauna is paramount for its conservation within and monitoring outside the native region. Here, we assemble a DNA barcode reference library for nearly 60% of all known Ponto-Caspian amphipod species. We use several methods to define molecular operational taxonomic units (MOTUs) based on two mitochondrial markers (COI and 16S) and assess their congruence with current species-level taxonomy based on morphology. Depending on the method, we find that 54–69% of species had congruent morpho-molecular boundaries. The cases of incongruence resulted from lumping distinct morphospecies into a single MOTU (7–27%), splitting a morphospecies into several MOTUs (4–28%), or both (4–11%). MOTUs defined by distance-based methods without a priori divergence thresholds showed the highest congruence with morphological taxonomy. These results indicate that DNA barcoding is valuable for clarifying the diversity of Ponto-Caspian amphipods, but reveals that extensive work is needed to resolve taxonomic uncertainties. Our study advances the DNA barcode reference library for the European aquatic biota, paving the way towards improved taxonomic knowledge needed to enhance monitoring and conservation efforts.

Climate change and fisheries exploitation are dramatically changing the species composition, abundances, and size spectra of fish communities. We explore whether variation in abundance-size spectra, a widely studied ecosystem feature, is influenced by a critical parameter thought to govern the shape of size-structured ecosystems—the relationship between the sizes of predators and their prey (predator-prey mass ratios, or PPMRs). PPMR estimates are lacking for vast numbers of fish species, including at the broader trophic guild scale. Using measurements of 8,128 prey items in gut contents of 97 reef fish species, we established PPMRs for four major trophic guilds (piscivores, invertivores, planktivores and herbivores) using linear mixed effects models. To assess theoretical predictions that higher mean community-level PPMR leads to shallower size spectrum slopes, we compared observations of mean community-level PPMR with size spectrum slopes for coastal reef sites distributed around Australia. PPMRs of individual fishes were remarkably high (median ~71,000), with significant variation between different trophic guilds (~890 for piscivores; ~83,000 for planktivores), and ~8,700 for whole communities. Community-level PPMRs were positively related to size spectrum slopes, broadly consistent with theory, however, this pattern was also influenced by the latitudinal temperature gradient. Tropical reefs showed a stronger relationship between community-level PPMRs and community size spectrum slopes than temperate reefs. The extent that these patterns apply outside Australia, and consequences for community structure and dynamics, are key areas for future investigation.

Resolving the combined effect of climate warming and exploitation in a food web context is key for predicting future biomass production, size-structure, and potential yields of marine fishes. Previous studies based on mechanistic size-based food web models have found that bottom-up processes are important drivers of size-structure and fisheries yield in changing climates. However, we know less about the joint effects of ‘bottom-up’ and ‘top-down’ effects of temperature: how do temperature effects propagate from individual-level physiology through food webs and alter the size-structure of exploited species in a community? Here we assess how a species-resolved size-based food web is affected by warming through both these pathways, and by exploitation. We parameterize a dynamic size spectrum food web model inspired by the offshore Baltic Sea food web, and investigate how individual growth rates, size-structure, relative abundances of species and yields are affected by warming. The magnitude of warming is based on projections by the regional coupled model system RCA4-NEMO and the RCP 8.5 emission scenario, and we evaluate different scenarios of temperature dependence on fish physiology and resource productivity. When accounting for temperature-effects on physiology in addition to on basal productivity, projected size-at-age in 2050 increases on average for all fish species, mainly for young fish, compared to scenarios without warming. In contrast, size-at-age decreases when temperature affects resource dynamics only, and the decline is largest for young fish. Faster growth rates due to warming, however, do not always translate to larger yields, as lower resource carrying capacities with increasing temperature tend to result in declines in the abundance of larger fish and hence spawning stock biomass – the part of the population exposed to fishing. These results show that to understand how global warming impacts the size structure of fish communities, both direct metabolic effects and indirect effects of temperature via basal resources must be accounted for.

As the popularity of recreational fishing gathers global momentum, so does the importance of knowing the number of active anglers and their spatial behaviour. Conventional counting methods, however, can be inaccurate and time-consuming. Here we present two novel methods to monitor recreational fishing applied in Kaunas water reservoir (ca 65 km2), Lithuania, comparing their performance to a conventional visual count. First, we employed a remotely piloted fixed wing drone which conducted 39 missions distributed over one year and compared its accuracy to conventional visual land or boat-based counts. With these data we developed a linear model to predict the annual number of anglers depending on weekday and ice conditions. Second, we used anonymous data from a popular GPS-enabled sonar device Deeper®, used by anglers to explore underwater landscapes and to find fish. The sonar usage probability was calibrated with angler observations from drones using Bayesian methods, demonstrating that at any given time ~2 % of anglers are using the sonar device during the open water season and ~15 % during the ice fishing season. The calibrated values were then used to estimate the total number of anglers, given the daily records of sonar usage in Kaunas water reservoir. The predicted annual number of anglers from both linear drone-based and Bayesian sonar-based methods gave similar results of 25 and 27 thousand anglers within the area during the period of day surveyed, which corresponded to nearly 110 thousand angling trips in the total reservoir area annually. Our study shows high potential of both drone and fish finder digital devices for assessing recreational fishing activities through space and time.

Many inland ecosystems (lakes, rivers, reservoirs, lagoons) around the world undergo regular biological monitoring surveys, including monitoring the abundance, biomass and size structure of fish communities. Yet, the majority of fish monitoring datasets for inland ecosystems remain inaccessible. This is especially true for historical datasets from the early and middle 20th century, despite their immense importance for establishing baselines of ecosystem status (e.g., prior to manifestations of climate change and intensive fisheries impacts), assessing the current status of fish stocks, and more generally determining temporal changes in fish populations. Here we present a newly digitized fish monitoring dataset for two major Lithuanian inland ecosystems – Curonian Lagoon and Kaunas Water Reservoir. The data comprises >60000 records from >800 fish surveys conducted during 1950s to 1980s, using a range of fishing gears and sampling methods. We introduce three different definitions for survey methods to describe the level of detail for each fish community study. Method 1 surveys include individual fish sizes and weights, Method 2 surveys record frequencies of fish in length or weight groups, whereas Method 3 only records the total catch biomass of a given species. The majority of historical and currently collected fish survey data can be attributed to one of these three methods and we present R codes to convert data from higher resolution methods into aggregated data formats, to facilitate data sharing. In addition, commercial fisheries catch data for years that were surveyed are also provided. The data presented here can facilitate ecological and fisheries analyses of baseline ecosystem status before the onsets of rapid warming and eutrophication, exploration of fish size structure, evaluation of different catch per unit effort standardization methods, and assessment of population responses to commercial fishing.

Marine ecosystem management is increasingly expected to take into account a wide range of ecological and socio‐economic factors. Decision‐making is helped by end‐to‐end ecosystem models that allow exploration of alternative management scenarios given a complex range of interacting factors. We present Atlantis – a spatially structured largely deterministic end‐to‐end marine ecosystem model written in C, available for all major operating systems, based on dynamically interacting physics, biology, fisheries, management, assessment and economics submodels. A detailed installation guide and example application files are also provided. One of the main features of Atlantis is its modularity. At the simplest level Atlantis can have uniform forcing of oceanographic processes, a single primary producer and a consumer. At the most complex level, Atlantis can be used with a range of environmentally driven ecological responses, complex and habitat‐dependent food web, dynamic assessment, management and fishing effort driven by market forces and human behaviour. The combination chosen should be guided by the available data and the questions to be answered. Atlantis provides a large and customizable list of output files and summary statistics that can be analysed and plotted using a number of dedicated r packages. When applying the Atlantis package, the users should be aware of the caveats associated with complex models, such as parameter and structural model uncertainty and challenges interpreting interactions of multiple processes.

Global warming and fisheries harvest are significantly impacting wild fish stocks, yet their interactive influence on population resilience to stress remains unclear. We explored these interactive effects on early-life development and survival by experimentally manipulating the thermal and harvest regimes in 18 zebrafish (Danio rerio) populations over six consecutive generations. Warming advanced development rates across generations, but after three generations, it caused a sudden and large (30–50%) decline in recruitment. This warming impact was most severe in populations where size-selective harvesting reduced the average size of spawners. We then explored whether our observed recruitment decline could be explained by changes in egg size, early egg and larval survival, population sex ratio, and developmental costs. We found that it was most likely driven by temperature-induced shifts in embryonic development rate and fishing-induced male-biased sex ratios. Importantly, once harvest and warming were relaxed, recruitment rates rapidly recovered. Our study suggests that the effects of warming and fishing could have strong impacts on wild stock recruitment, but this may take several generations to manifest. However, resilience of wild populations may be higher if fishing preserves sufficient body size diversity, and windows of suitable temperature periodically occur.

One of the most pronounced effects of climate change on the world’s oceans is the (generally) poleward movement of species and fishery stocks in response to increasing water temperatures. In some regions, such redistributions are already causing dramatic shifts in marine socioecological systems, profoundly altering ecosystem structure and function, challenging domestic and international fisheries, and impacting on human communities. Such effects are expected to become increasingly widespread as waters continue to warm and species ranges continue to shift. Actions taken over the coming decade (2021–2030) can help us adapt to species redistributions and minimise negative impacts on ecosystems and human communities, achieving a more sustainable future in the face of ecosystem change. We describe key drivers related to climate-driven species redistributions that are likely to have a high impact and influence on whether a sustainable future is achievable by 2030. We posit two different futures—a ‘business as usual’ future and a technically achievable and more sustainable future, aligned with the Sustainable Development Goals. We then identify concrete actions that provide a pathway towards the more sustainable 2030 and that acknowledge and include Indigenous perspectives. Achieving this sustainable future will depend on improved monitoring and detection, and on adaptive, cooperative management to proactively respond to the challenge of species redistribution. We synthesise examples of such actions as the basis of a strategic approach to tackle this global-scale challenge for the benefit of humanity and ecosystems. Supplementary Information The online version contains supplementary material available at 10.1007/s11160-021-09641-3.

Aim The negative correlation between temperature and body size of ectothermic animals (broadly known as the temperature-size rule or TSR) is a widely observed pattern, especially in aquatic organisms. Studies have claimed that TSR arises due to decreased oxygen solubility and increasing metabolic costs at warmer temperatures, whereby oxygen supply to a large body becomes increasingly difficult. However, mixed empirical evidence has led to a controversy about the mechanisms affecting species’ size and performance under different temperatures. We review the main competing genetic, physiological and ecological explanations for TSR and suggest a roadmap to move the field forward. Location Global Taxa Aquatic ectotherms Time period 1980 – Present Results We show that current studies cannot discriminate among alternative hypotheses and none of the hypotheses can explain all TSR-related observations. To resolve the impasse we need experiments and field-sampling programs that specifically compare alternative mechanisms and formally consider energetics related to growth costs, oxygen supply and behaviour. We highlight the distinction between evolutionary and plastic mechanisms, and suggest that the oxygen limitation debate should separate processes operating on short, decadal and millennial timescales. Conclusions Despite decades of research, we remain uncertain whether TSR is an adaptive response to temperature-related physiological (enzyme activity) or ecological changes (food, predation, other mortality), or a response to constraints operating at a cellular level (oxygen supply and associated costs). To make progress, ecologists, physiologists, modellers and geneticists should work together to develop a cross-disciplinary research program that integrates theory and data, explores time scales over which TSR operates, and assesses limits to adaptation or plasticity. We identify four questions for such a program. Answering these questions is crucial given the widespread impacts of climate change and reliance of management on models that are highly dependent on accurate representation of ecological and physiological responses to temperature. Keywords: adaptation, alternative mechanisms, climate change, growth, poikilotherm, energy budget, geometric biology, temperature size rule

Trade-offs in energy allocation between growth, reproduction, and survival are at the core of life-history theory. While age-specific mortality is considered to be the main determinant of the optimal allocation, some life-history strategies, such as delayed or skipped reproduction, may be better understood when also accounting for reproduction costs. Here, we present a two-pool indeterminate grower model that includes survival and energetic costs of reproduction. The energetic cost sets a minimum reserve required for reproduction, while the survival cost reflects increased mortality from low postreproductive body condition. Three life-history parameters determining age-dependent energy allocation to soma, reserve, and reproduction are optimized, and we show that the optimal strategies can reproduce realistic emergent growth trajectories, maturation ages, and reproductive outputs for fish. The model predicts maturation phase shifts along the gradient of condition-related mortality and shows that increased harvesting will select for earlier maturation and higher energy allocation to reproduction. However, since the energetic reproduction cost sets limits on how early an individual can mature, an increase in fitness at high harvesting can only be achieved by diverting most reserves into reproduction. The model presented here can improve predictions of life-history responses to environmental change and human impacts because key life-history traits such as maturation age and size, maximum body size, and size-specific fecundity emerge dynamically.

Identifying the origins and colonization routes of alien species is critical to understanding the mechanisms, consequences and management of invasions. This study aims to demonstrate and explain conflicting genetic signals in a widespread invasive Ponto-Caspian mysid Paramysis lacustris using mitochondrial DNA and microsatellite markers. Native Black Sea populations of P. lacustris are divided into three distinct phylogeographic groups: Danube–Dniester region, coastal Danube lakes, and Dnieper River delta. In the 1950s–1970s the species has been extensively used for relatively well documented transplantations in Eastern Europe. However, the populations in the newly established distribution area do not match any of the known native groups, and their genetic signal does not correspond with the recorded transplantation history. Using Bayesian Approximate Computation we explore alternative scenarios that could explain the unexpected mitochondrial and nuclear signals in the introduced populations. We suggest that reservoir construction and break-up of natural dispersal barriers on the River Dnieper have brought about a novel stock, probably admixed from a previously isolated and unknown “relict” population in the middle Dnieper and the genetically distinct Dnieper delta lineage. Through further intentional transfers this new P. lacustris lineage is now widely present in Eastern Europe and is likely to have new gene and trait combinations. The conflicting mitochondrial-nuclear signal in non-native populations also highlights the importance of using multiple genetic markers when tracing invasion sources and pathways.

Ectotherms generally shrink under experimental warming, but whether this pattern extends to wild populations is uncertain. We analysed ten million visual survey records, spanning the Australian continent and multiple decades and comprising the most common coastal reef fishes (335 species). We found that temperature indeed drives spatial and temporal changes in fish body size, but not consistently in the negative fashion expected. Around 55% of species were smaller in warmer waters (especially among small-bodied species), while 45% were bigger. The direction of a species' response to temperature through space was generally consistent with its response to temperature increase through time at any given location, suggesting that spatial trends could help forecast fish responses to long-term warming. However, temporal changes were about ten times faster than spatial trends (~4% versus ~40% body size change per 1 °C change through space and time, respectively). The rapid and variable responses of fish size to warming may herald unexpected impacts on ecosystem restructuring, with potentially greater consequences than if all species were shrinking.

Large vesicomyid clams (Veneroida: Vesicomyidae: Pliocardiinae) are prominent members of the communities associated with sulphide-rich deep-sea habitats. Taxonomic uncertainties within the Pliocardiinae result from both plasticity in shell morphologies and the common occurrence of cryptic species. Molecular taxonomic studies have now clarified many species-level assignments and provided DNA-barcodes for more than 50 species worldwide. Nonetheless, genus-level assignments remain uncertain, because the existing COI barcode sequences are not sufficient for identifying higher-level groupings. To construct a robust phylogeny for this subfamily, we conducted a combined Bayesian analysis of the COI mitochondrial fragment and five additional independent nuclear gene segments. The phylogenetic results provide a better foundation for assessing genus-level assignments within the subfamily and reveal goals for future taxonomic research. Furthermore, morphological examinations helped to clarify and solidify generic cl...

Evidence is accumulating that many marine ectotherms are undergoing rapid changes in their life-history characteristics. These changes have been variously attributed to fisheries-induced evolution, inhibited adult growth rate due to oxygen limitation at higher temperatures, and plastic responses to density dependence or changes in ocean productivity. Here, we review the diverse underlying mechanisms by which plastic and evolutionary responses to climate change and fisheries are likely to produce similar life-history trends in harvested marine ectotherms, leading to faster life-histories with earlier maturation and smaller adult size-at-age. While mechanistically understanding these growth and maturation changes may be difficult, it is becoming clear that changing life-histories will lead to modified population dynamics, productivity and natural mortality of the affected species. We discuss how the observed and expected life-history changes could affect the assumptions and uncertainty within single and multispecies models currently used in marine ecosystem management, highlighting that models which allow for dynamic life-history traits often report significantly different estimates of stock biomass. Given that both climate- and harvest-induced life-history changes are likely to intensify and possibly amplify each other, there is an urgent need to adequately assess the implications of faster life-histories for marine ecosystem management. This is especially true for data-poor stocks, where growth and maturation are not regularly assessed. Targeted monitoring can be used to inform responsive management, but for improved sustainability outcomes, a precautionary approach to management that is robust to life-history trends is advised.

Factors affecting population recovery from depletion are at the focus of wildlife management. Particularly, it has been debated how life-history characteristics might affect population recovery ability and productivity. Many exploited fish stocks have shown temporal changes towards earlier maturation and reduced adult body size, potentially owing to evolutionary responses to fishing. Whereas such life-history changes have been widely documented, their potential role on stock's ability to recover from exploitation often remains ignored by traditional fisheries management. We used a marine ecosystem model parameterized for Southeastern Australian ecosystem to explore how changes towards "faster" life histories might affect population per capita growth rate r. We show that for most species changes towards earlier maturation during fishing have a negative effect (3-40% decrease) on r during the recovery phase. Faster juvenile growth and earlier maturation were beneficial early in life, but smaller adult body sizes reduced the lifetime reproductive output and increased adult natural mortality. However, both at intra- and inter-specific level natural mortality and trophic position of the species were as important in determining r as species longevity and age of maturation, suggesting that r cannot be predicted from life-history traits alone. Our study highlights that factors affecting population recovery ability and productivity should be explored in a multi-species context, where both age-specific fecundity and survival schedules are addressed simultaneously. It also suggests that contemporary life-history changes in harvested species are unlikely to increase their resilience and recovery ability.

The Ponto-Caspian brackish-water fauna inhabits estuaries and rivers of the Black, Azov and Caspian seas and is fragmented by higher salinity waters and a major interbasin watershed. The fauna is known for the high levels of endemism, complex zoogeographic histories, and as a recent source of successful invasive species. It remains debated whether the Black and Azov Sea brackish-water populations survived unfavourable Pleistocene conditions in multiple separate refugia or whether the two seas were (repeatedly) recolonized from the Caspian. Using microsatellite and mtDNA markers, we demonstrate deep among- and within-basin subdivisions in a widespread Ponto-Caspian mysid crustacean Paramysis lacustris. Five genetic clusters were identified, but their relationships did not reflect the geography of the region. The Azov cluster was the most distinct (4-5% COI divergence), despite its geographic position in the corridor between Black and Caspian seas, and may represent a new species. In the northern Black Sea area, the Dnieper cluster was closer to the Caspian cluster than to the neighbouring Danube-Dniester-Bug populations, suggesting separate colonizations of the Black Sea. Overall, the data implied a predominant gene flow from the east to the Black Sea and highlight the importance of Caspian Sea transgressions in facilitating dispersal. Yet, the presence of distinct lineages in the Black Sea points to the persistence of isolated refugial populations that have gained diagnostic differences under presumably high mutation rates and large population sizes. The unfavourable Pleistocene periods in the Black Sea therefore appear to have promoted diversification of the brackish-water lineages, rather than extirpated them.

Many fished stocks show long-term reductions in adult body size. Such changes could lead to new feeding interactions and alter stock productivity, introducing new levels of uncertainty in fisheries management. We use a marine ecosystem model parameterized for Southeast Australia to explore how reductions (up to 6% in 50 years) in size-at-age of fished species affect stock recovery after an implementation of a fishing moratorium. We show that reduction in body size can greatly elevate predation mortality and lower the post-fishing biomass of affected species. In our simulations, the recovery period after the fishing moratorium was characterized by two phases. In the initial readjustment phase, the ecosystem dynamics was largely determined by the rapid changes in the biomasses of recovering species and changes in body size had negligible effects. In contrast, fish body sizes had the major impact on the biomasses in the second, semi-equilibrium state and the final biomasses were generally not affected by the harvest rate during the fishing period. When reduced size-at-age elevated predation mortality in most age groups of a species (tiger flathead Platycephalus richardsoni or silver warehou Seriolella punctata in our simulations), the species' equilibrium biomass was considerably lower compared with the scenarios of no change in body size. For other species (pink ling Genypterus blacodes and jackass morwong Nemadactylus macropterus), a predation increase in some age groups was balanced by the decrease in others. The latter reduction in predation mortality occurred when major predators of species with reducing size-at-age were decreasing in size themselves, or when cannibalism was an important source of juvenile mortality (in blue grenadier Macruronus novaezelandiae). We suggest that decreased size-at-age will be most detrimental to stock recovery when the main predators of the stock are not affected by the drivers causing changes in body size.

Calibration of complex, process-based ecosystem models is a timely task with modellers challenged by many parameters, multiple outputs of interest and often a scarcity of empirical data. Incorrect calibration can lead to unrealistic ecological and socio-economic predictions with the modeller’s experience and available knowledge of the modelled system largely determining the success of model calibration. Here we provide an overview of best practices when calibrating an Atlantis marine ecosystem model, a widely adopted framework that includes the parameters and processes comprised in many different ecosystem models. We highlight the importance of understanding the model structure and data sources of the modelled system. We then focus on several model outputs (biomass trajectories, age distributions, condition at age, realised diet proportions, and spatial maps) and describe diagnostic routines that can assist modellers to identify likely erroneous parameter values. We detail strategies to fine tune values of four groups of core parameters: growth, predator-prey interactions, recruitment and mortality. Additionally, we provide a pedigree routine to evaluate the uncertainty of an Atlantis ecosystem model based on data sources used. Describing best and current practices will better equip future modellers of complex, processed-based ecosystem models to provide a more reliable means of explaining and predicting the dynamics of marine ecosystems. Moreover, it promotes greater transparency between modellers and end-users, including resource managers.

Recent studies suggest that fishing and climate change can lead to a decrease in body size of fishes. While the effect of fishing on marine ecosystems has been widely explored, much less is known about the ecosystem effects of fish body size decrease. Here we used a marine ecosystem model to compare how fishing and small (

Large vesicomyid clams are common inhabitants of sulphidic deep-sea habitats such as hydrothermal vents, hydrocarbon seeps and whale-falls. Yet, the species- and genus-level taxonomy of these diverse clams has been unstable due to insufficiencies in sampling and absence of detailed taxonomic studies that would consistently compare molecular and morphological characters. To clarify uncertainties about species-level assignments, we examined DNA sequences from mitochondrial cytochrome-c-oxidase subunit I (COI) in conjunction with morphological characters. New and published COI sequences were used to create a molecular database for 44 unique evolutionary lineages corresponding to species. Overall, the congruence between molecular and morphological characters was good. Several discrepancies due to synonymous species designations were recognized, and acceptable species names were rectified with published COI sequences in cases where morphological specimens were available. We identified seven species with trans-...

The hypothesis that selection on the opsin gene is efficient in tuning vision to the ambient light environment of an organism was assessed in 49 populations of 12 Mysis crustacean species, inhabiting arctic marine waters, coastal littoral habitats, freshwater lakes (‘glacial relicts’) and the deep Caspian Sea. Extensive sequence variation was found within and among taxa, but its patterns did not match expectations based on light environments, spectral sensitivity of the visual pigment measured by microspectrophotometry or the history of species and populations. The main split in the opsin gene tree was between lineages I and II, differing in six amino acids. Lineage I was present in marine and Caspian Sea species and in the North American freshwater Mysis diluviana, whereas lineage II was found in the European and circumarctic fresh- and brackish-water Mysis relicta, Mysis salemaai and Mysis segerstralei. Both lineages were present in some populations of M. salemaai and M. segerstralei. Absorbance spectra of the visual pigment in nine populations of the latter three species showed a dichotomy between lake (λmax = 554–562 nm) and brackish-water (Baltic Sea) populations (λmax = 521–535 nm). Judged by the shape of spectra, this difference was not because of different chromophores (A2 vs. A1), but neither did it coincide with the split in the opsin tree (lineages I/II), species identity or current light environments. In all, adaptive evolution of the opsin gene in Mysis could not be demonstrated, but its sequence variation did not conform to a neutral expectation either, suggesting evolutionary constraints and/or unidentified mechanisms of spectral tuning.

The study reports new primers capable of amplifying fragments from three nuclear protein-coding genes in a variety of deep-sea molluscs and annelids - adenine nucleotide translocase (Ant), calmodulin (Cal) and cyclophilin A (CycA). The Ant primers appear to be restricted to bivalve molluscs, whereas the Cal and CycA primers also amplified appropriate gene fragments from Lepetodrilus gastropod molluscs and Osedax polychaete worms. The amplified fragment of Cal contains an intron in the molluscs, but no intron was detected in the Ant and CycA fragments from any of the tested animals. DNA sequences generated by the three primer sets exhibited one to 15 single nucleotide polymorphism sites in deep-sea vesicomyid clams and Osedax boneworms. The observed levels of polymorphism indicate that the genes are likely to be useful in both population genetic and phylogenetic analyses of different invertebrate taxa.

Aim Limnomysis benedeni Czerniavsky, 1882 is a mysid crustacean native to the Ponto-Caspian (Black and Caspian Sea) rivers and estuaries, and has recently spread across Europe through intentional and unintentional introductions. We explored the structuring of genetic variation in native and non-native populations with an aim to trace the sources of the invasions, and to infer whether the spread has occurred through a single or multiple invasion waves. Location Native estuaries in the Ponto-Caspian basin (Volga, Don, Dnieper, Dniester, Danube) and the recently colonized range along the Danube–Rhine river systems and Lithuania. Methods A fragment of the mitochondrial COI gene was sequenced to assess genetic affinities and diversity in native and recently established populations. Results The genetic diversity in the native regions is organized into several strongly diverged haplotype groups or lineages, partly allopatric, partly sympatric. All these lineages have also spread beyond the native range. Even the recent rapid dispersal across Europe along the Danube–Rhine system towards the North Sea basin involved several lineages from the Danube delta sector. The structuring of genetic diversity among invaded sites suggests multiple invasion events to the Danube–Rhine drainage. This contrasts with data from some other Ponto-Caspian species, where a single haplotype seems to have occupied most invaded areas. There is no evidence that intentionally stocked reservoirs in the Baltic Sea basin would have contributed to further unintentional spread of L. benedeni. Main conclusions Limnomysis benedeni is spreading across Europe using the southern invasion corridor. The invasion most likely involved several waves from differentiated sources in the native Danube delta area.

A presence of two cryptic biological species of Hediste diversicolor complex polychaetes was corroborated in a geographical survey of some 30 populations from the eastern and southern coasts of the Baltic Sea, with data from four completely diagnostic allozyme characters. Species A was dominant in the northernmost part of the Baltic Hediste range (Bothnian Sea), whereas Species B alone was found in the south (Poland, Germany, Denmark). In the intervening region, comprising the majority of the sites studied in southern Finland and Estonia, the two species were usually found together, with no evidence of recent hybridisation (i.e., no heterozygote genotypes). While mitochondrial DNA also distinguished the two taxa, it was not similarly completely diagnostic, but there were rare cases (ca 5%) of lineage mismatch indicating that some introgression has occurred in the past. Comparison with published data suggests that species A also inhabits the North Sea–NE Atlantic–Mediterranean coasts, and species B is also present in the North Sea and the NW Atlantic (Canada). Within the Baltic, the two species show distinctly different patterns of mtDNA diversity, plausibly related to different colonisation histories. Species A shows a generally high haplotype and nucleotide diversity, whereas in species B we found only four deeply diverged groups of closely related haplotypes. Hypothetically this could indicate a recent expansion of species B from a small number of colonising individuals. Moreover, species B showed marked intraspecific geographical structuring, with co-incident genetic changes along the N Estonian–S Finnish coasts both in mtDNA and an allozyme marker; this pattern suggests a contact between two genetically distinct invasion waves of different origins. In all, species A and B represent good, reproductively isolated and partly sympatric species which require to be recognised in ecological work. A formal taxonomical description is needed, but awaits better, range-wide distributional and ecological characterisation and working out of morphological differences that enable a practical identification.

The Ponto-Caspian (Black and Caspian seas) brackish-water fauna represents a special case of the endemic diversification in world's ancient lakes; it also involves a hotspot of continental diversity in the predominantly marine mysid crustaceans. We explored the origins and history of the mysid diversification in a phylogenetic analysis of some 20 endemic Ponto-Caspian species mainly of the genus Paramysis and their marine congeners, using sequences of two nuclear protein-coding genes, two nuclear rRNA genes, the mitochondrial COI gene and morphological data. A nearly completely resolved phylogeny was recovered, with no indication of rapid diversification bursts. Deep divergences were found among the main endemic clades, attesting to a long independent faunal history in the continental Paratethys waters. The current marine Paramysis species make a monophyletic cluster secondarily derived from the continental Paratethyan (Ponto-Caspian) Paramysis ancestors. The good phylogenetic resolution was mainly due to the two nuclear protein-coding genes, opsin and EPRS, here for the first time applied to peracarid systematics. In contrast, 'conventional' mtDNA and nuclear rRNA genes provided poor topological resolution and weak congruence of divergence rates. The two nuclear protein-coding genes had more congruent rates of evolution, and were about 10-15 times slower than the mitochondrial COI gene.

The Ponto-Caspian mysid crustacean Paramysis bakuensis G.O. Sars, 1895, which was previously synonymized with P. baeri Czerniavsky, 1882, is restored on the basis of new morphological and molecular characters. The Sea of Azov subspecies P. baeri bispinosa Martynov, 1924, in turn, is synonymised with P. bakuensis. The two species, P. baeri and P. bakuensis, are distinguished by the shapes of paradactylar setae of pereiopods, maxilla II exopod and antennal scale, and by the number of denticles in the telson cleft. They also are characterized by ca 7% divergence in mitochondrial COI gene sequences. P. bakuensis is shown to be a widespread species, distributed in estuaries and rivers of the Caspian, Azov and Black Sea basins and in the Caspian Sea itself. P. baeri is endemic to the Caspian Sea, where the two species overlap and are sometimes found together.