Invasion of the four kingdoms: the parasite journey across plant and non-plant hosts
Parasites have a rich and long natural history among biological entities, and it has been suggested that parasites are one of the most significant factors in the evolution of their hosts. However, it has been emphasized less frequently how co-evolution has undoubtedly also shaped the paths of parasites. It may seem safe to assume that specific differences among the array of potential hosts for particular parasites have restricted and diversified their evolutionary pathways and strategies for survival. Nevertheless, if one looks closely enough at host and parasite, one finds commonalities, both in terms of host defences and parasite strategies to out-manoeuvre them. While such analyses have been the source of numerous reviews, they are generally limited to interactions between, at most, one kingdom of parasite with two kingdoms of host (e.g. similarities in animal and plant host responses against fungi). With the aim of extending this view, we herein critically evaluate the similarities and differences across all four eukaryotic host kingdoms (plants, animals, fungi, and protists) and their parasites. In doing so, we show that hosts tend to share common strategies for defence, including both physical and behavioural barriers, and highly evolved immune responses, in particular innate immunity. Parasites have, similarly, evolved convergent strategies to counter these defences, including mechanisms of active penetration, and evading the host's innate and/or adaptive immune responses. Moreover, just as hosts have evolved behaviours to avoid parasites, many parasites have adaptations to manipulate host phenotype, physiologically, reproductively, and in terms of behaviour. Many of these strategies overlap in the host and parasite, even across wide phylogenetic expanses. That said, specific differences in host physiology and immune responses often necessitate different adaptations for parasites exploiting fundamentally different hosts. Taken together, this review facilitates hypothesis-driven investigations of parasite-host interactions that transcend the traditional kingdom-based research fields.
Molluscan systematics: historical perspectives and the way ahead
Mollusca, the second-most diverse animal phylum, is estimated to have over 100,000 living species with great genetic and phenotypic diversity, a rich fossil record, and a considerable evolutionary significance. Early work on molluscan systematics was grounded in morphological and anatomical studies. With the transition from oligo gene Sanger sequencing to cutting-edge genomic sequencing technologies, molecular data has been increasingly utilised, providing abundant information for reconstructing the molluscan phylogenetic tree. However, relationships among and within most major lineages of Mollusca have long been contentious, often due to limited genetic markers, insufficient taxon sampling and phylogenetic conflict. Fortunately, remarkable progress in molluscan systematics has been made in recent years, which has shed light on how major molluscan groups have evolved. In this review of molluscan systematics, we first synthesise the current understanding of the molluscan Tree of Life at higher taxonomic levels. We then discuss how micromolluscs, which have adult individuals with a body size smaller than 5 mm, offer unique insights into Mollusca's vast diversity and deep phylogeny. Despite recent advancements, our knowledge of molluscan systematics and phylogeny still needs refinement. Further advancements in molluscan systematics will arise from integrating comprehensive data sets, including genome-scale data, exceptional fossils, and digital morphological data (including internal structures). Enhanced access to these data sets, combined with increased collaboration among morphologists, palaeontologists, evolutionary developmental biologists, and molecular phylogeneticists, will significantly advance this field.
The underlying causes of differential migration: assumptions, hypotheses, and predictions
Mechanisms governing the migratory decisions of birds have long fascinated ecologists and sparked considerable debate. Identifying factors responsible for variation in migration distance, also known as differential migration, has been a popular approach to understanding the mechanisms underlying migratory behaviour more generally. However, research progress has been slowed by the continued testing of overlapping, non-mechanistic, and circular predictions among a small set of historically entrenched hypotheses. We highlight the body size hypothesis and suggest that the predictions commonly tested have impeded progress because body size relationships with migration distance are predictions made by several distinct hypotheses with contrasting mechanisms. The cost of migration itself has not been adequately accounted for in most hypotheses, and we propose two flight efficiency hypotheses with time- and energy-minimizing mechanisms that allow individuals to mitigate the risks inherent to longer migrations. We also advance two conceptual versions of the social dominance hypothesis based on two distinct underlying mechanisms related to distance minimization and food maximization that will help clarify the role of competition in driving migratory decisions. Overall, we describe and refine 12 mechanistic hypotheses proposed to explain differential migration (along with several other special-case hypotheses), seven of which have underlying mechanisms related to food limitation as past research has identified this to be an important driver of differential migration. We also thoroughly reviewed 145 publications to assess the amount of support for 10 critical assumptions underlying alternative hypotheses for differential migration in birds. Our review reveals that surprisingly few studies explicitly evaluate assumptions within a differential migration context. Generating and testing strong predictions and critical assumptions underlying mechanisms of alternative hypotheses will improve our ability to differentiate among these explanations of differential migration. Additionally, future intraspecific progress will be greatest if investigators continue to focus on mechanisms underlying variation in migration distance within rather than among demographic classes, as previous research has found differing mechanisms to be responsible for differential migration among demographic classes. Interspecifically, a thorough comparative analysis that seeks to explain variation in migration distance among species would broaden both our understanding of the mechanisms regulating current differential migration patterns and those that led to the evolution of migration more generally. Collectively, we provide a framework that, together with advances in animal-borne tracking and other technology, can be used to advance our understanding of the causes of differential migration distance, and migratory decisions more generally.
Development of eodiscinid trilobites
A comprehensive review of a full developmental sequence of eodiscinid trilobites reported in recent decades from Cambrian Series 2 and 3 strata is presented. These mostly articulated specimens exhibit detailed morphologies with preservation of even delicate structures at different growth stages, such as the bacculae, axial pores, spine pores, pygidial marginal spines and line coaptative structures. Their trunk segmentation schedules displayed a consistent developmental mode in segment generation and liberation, that is tagmosis and somitogenesis occurred heterochronously after each moulting event, providing clues regarding the potential developmental strategy in isopygous and even macropygous trilobites. The fact that the rate of segmentation obviously exceeds that of articulation results in a seemingly prolonged process of the formation of thoracic segments, which might explain why eodiscinid trilobites have a fixed and limited number of thoracic segments. In addition, the relationship between enrollment mechanism and trunk segmentation during eodiscinid ontogeny confirms this highly unusual growth pattern among the Trilobita, revealing why these early-diverging trilobites controlled the rate of segment increase and release during their life cycles, and is thus of interest with regard to the evolution of arthropod body patterning.
Does death drive the scaling of life?
The magnitude of many kinds of biological structures and processes scale with organismal size, often in regular ways that can be described by power functions. Traditionally, many of these "biological scaling" relationships have been explained based on internal geometric, physical, and energetic constraints according to universal natural laws, such as the "surface law" and "3/4-power law". However, during the last three decades it has become increasingly apparent that biological scaling relationships vary greatly in response to various external (environmental) factors. In this review, I propose and provide several lines of evidence supporting a new ecological perspective that I call the "mortality theory of ecology" (MorTE). According to this viewpoint, mortality imposes time limits on the growth, development, and reproduction of organisms. Accordingly, small, vulnerable organisms subject to high mortality due to predation and other environmental hazards have evolved faster, shorter lives than larger, more protected organisms. A MorTE also includes various corollary, size-related internal and external causative factors (e.g. intraspecific resource competition, geometric surface area to volume effects on resource supply/transport and the protection of internal tissues from environmental hazards, internal homeostatic regulatory systems, incidence of pathogens and parasites, etc.) that impact the scaling of life. A mortality-centred approach successfully predicts the ranges of body-mass scaling slopes observed for many kinds of biological and ecological traits. Furthermore, I argue that mortality rate should be considered the ultimate (evolutionary) driver of the scaling of life, that is expressed in the context of other proximate (functional) drivers such as information-based biological regulation and spatial (geometric) and energetic (metabolic) constraints.
Comparative gut microbiome research through the lens of ecology: theoretical considerations and best practices
Comparative approaches in animal gut microbiome research have revealed patterns of phylosymbiosis, dietary and physiological convergences, and environment-host interactions. However, most large-scale comparative studies, especially those that are highly cited, have focused on mammals, and efforts to integrate comparative approaches with existing ecological frameworks are lacking. While mammals serve as useful model organisms, developing generalised principles of how animal gut microbiomes are shaped and how these microbiomes interact bidirectionally with host ecology and evolution requires a more complete sampling of the animal kingdom. Here, we provide an overview of what past comparative studies have taught us about the gut microbiome, and how community ecology theory may help resolve certain contradictions in comparative gut microbiome research. We explore whether certain hypotheses are supported across clades, and how the disproportionate focus on mammals has introduced potential bias into gut microbiome theory. We then introduce a methodological solution by which public gut microbiome data of understudied hosts can be compiled and analysed in a comparative context. Our aggregation and analysis of 179 studies shows that generating data sets with rich host diversity is possible with public data and that key gut microbes associated with mammals are widespread across the animal kingdom. We also show the effects that sample size and taxonomic rank have on comparative gut microbiome studies and that results of multivariate analyses can vary significantly with these two parameters. While challenges remain in developing a universal model of the animal gut microbiome, we show that existing ecological frameworks can help bring us one step closer to integrating the gut microbiome into animal ecology and evolution.
Archaeocytes in sponges: simple cells of complicated fate
Archaeocytes are considered a key cell type in sponges (Porifera). They are believed to be multifunctional cells performing various functions, from nutrient digestion to acting as adult stem cells (ASCs). Thus, archaeocytes are mentioned in discussions on various aspects of sponge biology. As presumed ASCs of an early-diverged animal taxon, archaeocytes are of great fundamental interest for further progress in understanding tissue functioning in metazoans. However, the term 'archaeocyte' is rather ambiguous in its usage and understanding, and debates surrounding archaeocytes have persisted for over a century, reflecting the ongoing complexity of understanding their nature. This article presents a comprehensive revision of the archaeocyte concept, including both its historical development and biological features (i.e. taxonomic distribution, characteristics, and functions). The term 'archaeocyte' and its central aspects were introduced as early as the end of the 19th century based on data mainly from demosponges. Remarkably, despite the general lack of comparative and non-histological data, these early studies already regarded archaeocytes as the ASCs of sponges. These early views were readily inherited by subsequent studies, often without proper verification, shaping views on many aspects of sponge biology for more than a century. Taking into account all available data, we propose considering the archaeocytes as a cell type specific to the class Demospongiae. Clear homologues of archaeocytes are absent in other sponge classes. In demosponges, the term 'archaeocytes' refers to mesohyl cells that have an amoeboid shape, nucleolated nuclei, and non-specific inclusions in the cytoplasm. The absence of specific traits makes the archaeocytes a loosely defined and probably heterogeneous cell population, rendering the exhaustive characterisation of the 'true' archaeocyte population impossible. At the same time, the molecular characterisation of archaeocytes is only beginning to develop. Stemness and almost unlimited potency have always been at the core of the traditional archaeocyte concept. However, currently, the most consistent data on archaeocyte stem cell function come only from developing gemmules of freshwater sponges. For tissues of adult demosponges, the data favour a two-component stem cell system, in which archaeocytes may cooperate with another stem cell population, choanocytes. Simultaneously, cells with archaeocyte morphology function as macrophages in demosponges, participating in the food digestion cycle and immune defence. Such cells should be denoted with the more neutral term 'nucleolar amoebocytes', as the term 'archaeocyte' not only describes the morphology of a cell but also introduces the proposition of its stem nature. Thus, the future usage of the term 'archaeocyte' should be limited to cases where a cell is shown or at least presumed to be a stem cell.
Meta-analysis of the acoustic adaptation hypothesis reveals no support for the effect of vegetation structure on acoustic signalling across terrestrial vertebrates
Acoustic communication plays a prominent role in various ecological and evolutionary processes involving social interactions. The properties of acoustic signals are thought to be influenced not only by the interaction between signaller and receiver but also by the acoustic characteristics of the environment through which the signal is transmitted. This conjecture forms the core of the so-called "acoustic adaptation hypothesis" (AAH), which posits that vegetation structure affects frequency and temporal parameters of acoustic signals emitted by a signaller as a function of their acoustic degradation properties. Specifically, animals in densely vegetated "closed habitats" are expected to produce longer acoustic signals with lower repetition rates and lower frequencies (minimum, mean, maximum, and peak) compared to those inhabiting less-vegetated "open habitats". To date, this hypothesis has received mixed results, with the level of support depending on the taxonomic group and the methodology used. We conducted a systematic literature search of empirical studies testing for an effect of vegetation structure on acoustic signalling and assessed the generality of the AAH using a meta-analytic approach based on 371 effect sizes from 75 studies and 57 taxa encompassing birds, mammals and amphibians. Overall, our results do not provide consistent support for the AAH, neither in within-species comparisons (suggesting no overall phenotypically plastic response of acoustic signalling to vegetation structure) nor in among-species comparisons (suggesting no overall evolutionary response). However, when considering birds only, we found weak support for the AAH in within-species comparisons, which was mainly driven by studies that measured frequency bandwidth, suggesting that this variable may exhibit a phenotypically plastic response to vegetation structure. For among-species comparisons in birds, we also found support for the AAH, but this effect was not significant after excluding comparative studies that did not account for phylogenetic non-independence. Collectively, our synthesis does not support a universal role of vegetation structure in the evolution of acoustic communication. We highlight the need for more empirical work on currently under-studied taxa such as amphibians, mammals, and insects. Furthermore, we propose a framework for future research on the AAH. We specifically advocate for a more detailed and quantitative characterisation of habitats to identify frequencies with the highest detection probability and to determine if frequencies with greater detection distances are preferentially used. Finally, we stress that empirical tests of the AAH should focus on signals that are selected for increased transmission distance.
Zooming in the plastisphere: the ecological interface for phytoplankton-plastic interactions in aquatic ecosystems
Phytoplankton is an essential resource in aquatic ecosystems, situated at the base of aquatic food webs. Plastic pollution can impact these organisms, potentially affecting the functioning of aquatic ecosystems. The interaction between plastics and phytoplankton is multifaceted: while microplastics can exert toxic effects on phytoplankton, plastics can also act as a substrate for colonisation. By reviewing the existing literature, this study aims to address pivotal questions concerning the intricate interplay among plastics and phytoplankton/phytobenthos and analyse impacts on fundamental ecosystem processes (e.g. primary production, nutrient cycling). This investigation spans both marine and freshwater ecosystems, examining diverse organisational levels from subcellular processes to entire ecosystems. The diverse chemical composition of plastics, along with their variable properties and role in forming the "plastisphere", underscores the complexity of their influences on aquatic environments. Morphological changes, alterations in metabolic processes, defence and stress responses, including homoaggregation and extracellular polysaccharide biosynthesis, represent adaptive strategies employed by phytoplankton to cope with plastic-induced stress. Plastics also serve as potential habitats for harmful algae and invasive species, thereby influencing biodiversity and environmental conditions. Processes affected by phytoplankton-plastic interaction can have cascading effects throughout the aquatic food web via altered bottom-up and top-down processes. This review emphasises that our understanding of how these multiple interactions compare in impact on natural processes is far from complete, and uncertainty persists regarding whether they drive significant alterations in ecological variables. A lack of comprehensive investigation poses a risk of overlooking fundamental aspects in addressing the environmental challenges associated with widespread plastic pollution.
Testosterone mediates life-history trade-offs in female mammals
Hormones mediate life-history trade-offs. In female mammals, such trade-offs have been studied predominantly in the contexts of oestrogen, progesterone and prolactin. We evaluate the hypothesis that prenatal and postnatal testosterone levels structure and regulate trade-offs in females involving components of reproduction and survival. This hypothesis is predicated on the observation that testosterone confers competition-related and survival-related benefits, but also reproduction-related costs, to female mammals. The hypothesis is supported by field and laboratory data from diverse non-human animals, and data from healthy women. Most broadly, relatively low testosterone level in females has been associated with earlier, faster and higher offspring production, greater attractiveness to males, and reduced dominance or competitiveness, whereas higher testosterone level is associated with delayed and reduced reproduction but increased dominance, status, aggression, and resource accrual. The magnitude of testosterone-mediated trade-offs is expected to depend upon the strength of female-female competition, which represents some function of species-specific ecology, behaviour and mating system. Testosterone-associated trade-offs have, until now, been virtually ignored in studies of female life history, reproductive physiology, evolutionary endocrinology, and female-limited disease, probably due to researcher biases towards conceptualizing androgens as hormones with effects mainly restricted to males.
Bringing together but staying apart: decisive differences in animal and fungal mitochondrial inner membrane fusion
Mitochondria are dynamic and plastic, undergoing continuous fission and fusion and rearrangement of their bioenergetic sub-compartments called cristae. These fascinating processes are best understood in animal and fungal models, which are taxonomically grouped together in the expansive Opisthokonta supergroup. In opisthokonts, crista remodelling and inner membrane fusion are linked by dynamin-related proteins (DRPs). Animal Opa1 (optical atrophy 1) and fungal Mgm1 (mitochondrial genome maintenance 1) are tacitly considered orthologs because their similar mitochondria-shaping roles are mediated by seemingly shared biochemical properties, and due to their presence in the two major opisthokontan subdivisions, Holozoa and Holomycota, respectively. However, molecular phylogenetics challenges this notion, suggesting that Opa1 and Mgm1 likely had separate, albeit convergent, evolutionary paths. Herein, we illuminate disparities in proteolytic processing, structure, and interaction network that may have bestowed on Opa1 and Mgm1 distinct mechanisms of membrane remodelling. A key disparity is that, unlike Mgm1, Opa1 directly recruits the mitochondrial phospholipid cardiolipin to remodel membranes. The differences outlined herein between the two DRPs could have broader impacts on mitochondrial morphogenesis. Outer and inner membrane fusion are autonomous in animals, which may have freed Opa1 to repurpose its intrinsic activity to remodel cristae, thereby regulating the formation of respiratory chain supercomplexes. More significantly, Opa1-mediated crista remodelling has emerged as an integral part of cytochrome c-regulated apoptosis in vertebrates, and perhaps in the cenancestor of animals. By contrast, outer and inner membrane fusion are coupled in budding yeast. Consequently, Mgm1 membrane-fusion activity is inextricable from its role in the biogenesis of fungal lamellar cristae. These disparate mitochondrial DRPs ultimately may have contributed to the different modes of multicellularity that have evolved within Opisthokonta.
Towards ecosystem-based techniques for tipping point detection
An ecosystem shifts to an alternative stable state when a threshold of accumulated pressure (i.e. direct impact of environmental change or human activities) is exceeded. Detecting this threshold in empirical data remains a challenge because ecosystems are governed by complex interlinkages and feedback loops between their components and pressures. In addition, multiple feedback mechanisms exist that can make an ecosystem resilient to state shifts. Therefore, unless a broad ecological perspective is used to detect state shifts, it remains questionable to what extent current detection methods really capture ecosystem state shifts and whether inferences made from smaller scale analyses can be implemented into ecosystem management. We reviewed the techniques currently used for retrospective detection of state shifts detection from empirical data. We show that most techniques are not suitable for taking a broad ecosystem perspective because approximately 85% do not combine intervariable non-linear relationships and high-dimensional data from multiple ecosystem variables, but rather tend to focus on one subsystem of the ecosystem. Thus, our perception of state shifts may be limited by methods that are often used on smaller data sets, unrepresentative of whole ecosystems. By reviewing the characteristics, advantages, and limitations of the current techniques, we identify methods that provide the potential to incorporate a broad ecosystem-based approach. We therefore provide perspectives into developing techniques better suited for detecting ecosystem state shifts that incorporate intervariable interactions and high-dimensionality data.
Insect-flower interactions, ecosystem functions, and restoration ecology in the northern Sahel: current knowledge and perspectives
Actions for ecological restoration under the Great Green Wall (GGW) initiative in the northern Sahel have been plant focused, paying scant attention to plant-animal interactions that are essential to ecosystem functioning. Calls to accelerate implementation of the GGW make it timely to develop a more solid conceptual foundation for restoration actions. As a step towards this goal, we review what is known in this region about an important class of plant-animal interactions, those between plants and flower-visiting insects. Essential for pollination, floral resources also support insects that play important roles in many other ecosystem processes. Extensive pastoralism is the principal subsistence mode in the region, and while recent analyses downplay the impact of livestock on vegetation dynamics compared to climatic factors, they focus primarily on rangeland productivity, neglecting biodiversity, which is critical for long-term sustainability. We summarise current knowledge on insect-flower interactions, identify information gaps, and suggest research priorities. Most insect-pollinated plants in the region have open-access flowers exploitable by diverse insects, an advantageous strategy in environments with low productivity and seasonal and highly variable rainfall. Other plant species have diverse traits that constrain the range of visitors, and several distinct flower types are represented, some of which have been postulated to match classical "pollination syndromes". As in most ecosystems, bees are among the most important pollinators. The bee fauna is dominated by ground-nesting solitary bees, almost all of which are polylectic. Many non-bee flower visitors also perform various ecosystem services such as decomposition and pest control. Many floral visitors occupy high trophic levels, and are indicators of continued functioning of the food webs on which they depend. The resilience of insect-flower networks in this region largely depends on trees, which flower year-round and are less affected by drought than forbs. However, the limited number of abundant tree species presents a potential fragility. Flowering failure of a crucial "hub" species during exceptionally dry years could jeopardise populations of some flower-visiting insects. Furthermore, across Sahelian drylands, browsers are increasingly predominant over grazers. Although better suited to changing climates, browsers exert more pressure on trees, potentially weakening insect-flower interaction networks. Understanding the separate and combined effects of climate change and land-use change on biotic interactions will be key to building a solid foundation to facilitate effective restoration of Sahelian ecosystems.
Insect immunity in the Anthropocene
Anthropogenic activities result in global change, including climate change, landscape degradation and pollution, that can alter insect physiology and immune defences. These changes may have contributed to global insect decline and the dynamics of insect-transmitted diseases. The ability of insects to mount immune responses upon infection is crucial for defence against pathogens and parasites. Suppressed immune defences reduce fitness by causing disease-driven mortality and elevated immune responses reduce energy available to invest in other fitness traits such as reproduction. Understanding the impact of anthropogenic factors on insect-pathogen interactions is therefore key to determining the contribution of anthropogenic global change to pathogen-driven global insect decline and the emergence and transmission of insect-borne diseases. Here, we synthesise evidence of the impact of anthropogenic factors on insect immunity. We found evidence that anthropogenic factors, such as insecticides and heavy metals, directly impacting insect immune responses by inhibiting immune activation pathways. Alternatively, factors such as global warming, heatwaves, elevated CO and landscape degradation can indirectly reduce insect immune responses via reducing the energy available for immune function. We further review how anthropogenic factors impact pathogen clearance and contribute to an increase in vector-borne diseases. We discuss the fitness cost of anthropogenic factors via pathogen-driven mortality and reduced reproductive output and how this can contribute to species extinction. We found that most research has determined the impact of a single anthropogenic factor on insect immune responses or pathogen resistance. We recommend studying the combined impact of multiple stressors on immune response and pathogen resistance to understand better how anthropogenic factors affect insect immunity. We conclude by highlighting the importance of initiatives to mitigate the impact of anthropogenic factors on insect immunity, to reduce the spread of vector-borne diseases, and to protect vulnerable ecosystems from emerging diseases.
Shedding light on biodiversity: reviewing existing knowledge and exploring hypothesised impacts of agrophotovoltaics
The growing demand for energy and the shift towards green energy solutions have led to the conversion of open spaces and agricultural fields into photovoltaic (PV) power plants, exacerbating the "food-energy-environment" trilemma. Agrophotovoltaics (APVs), a dual-use system combining agriculture and energy production on the same land, presents a potential solution to this challenge. While the environmental impacts of ground-mounted utility-scale PV (USPV) power plants and the effects of APV systems on agricultural yields have been extensively studied and reviewed, the implications for wildlife and biodiversity remain largely unexplored. This knowledge gap is pressing, given the accelerated global adoption of APV systems and the urgency of understanding their broader ecological consequences. In this concise review, we synthesise existing literature on the impacts of USPV installations on biodiversity and the effects of APV on crop production. Building on these foundations, we propose novel hypotheses concerning the potential pathways and mechanisms through which APV systems may influence biodiversity. We explore the complex interactions between agroecosystems and natural ecosystems, examining both direct and indirect effects. Our review culminates in a set of key research questions designed to guide future studies on the biodiversity outcomes of APV deployment. Future research should comprehensively address factors such as habitat type, climate, spatial scale, technology, and agricultural practices, as well as the overarching impacts of climate change. By highlighting the importance of these variables, we aim to facilitate a nuanced understanding of how APV systems can either support or undermine biodiversity. This work not only underscores the critical need for empirical studies in this emerging field but also sets the stage for more informed and sustainable implementation of APV technologies.
The ecological and evolutionary dynamics of inselbergs
Islands are fundamental model systems in ecology, biogeography, and evolutionary biology. However, terrestrial islands, unlike their aquatic counterparts, have received comparatively less attention. Among these land islands, inselbergs (i.e. isolated rock outcrops with diverse lithologies and a modest topographical prominence) stand out as iconic examples distributed worldwide across global biomes. Due to their durable lithology, inselbergs change slowly, persisting for tens of millions of years. In this review, we propose a biological definition for inselbergs that captures three fundamental characteristics of inselbergs from the perspective of biota. These are old age, isolation and the presence of unique microhabitats that are rare or absent in the surrounding matrix, fostering distinct communities often with unique and endemic biota. We synthesise the state of the art and formulate a set of testable hypotheses to deepen our understanding of the origins and maintenance of diversity on inselbergs, which are increasingly exposed to anthropogenic threats. By offering different habitats compared to the surrounding habitat matrix (e.g. moist microhabitats in dryland landscapes and xeric environments in humid tropical landscapes), inselbergs may allow specific lineages to thrive beyond their typical geographical limits. Particularly in drylands and degraded landscapes, inselbergs may not just provide different habitats but also act as ecological refuges or evolutionary refugia by providing a wider range of potential microhabitats than the surrounding matrix, enhancing resilience and promoting regional biodiversity. The central role of the matrix ensures that the ecological and evolutionary dynamics of inselbergs differ from those of true islands such as oceanic islands. Given that inselberg biota coexist within a terrestrial matrix, interactions between inselberg and matrix populations impact each other significantly. Over evolutionary timescales, matrix species may contract to inselberg refugia, preserving lineages while cycles of isolation and reconnection may drive speciation via a species pump. Although inselberg biodiversity has been studied predominantly from an island biogeography perspective, we argue that depending on the spatial scale, habitat specificity and mobility of the organisms considered, a range of different theories and paradigms can help explain the biogeography and local distribution patterns of different taxonomic and functional groups of inselberg species.
EthoCRED: a framework to guide reporting and evaluation of the relevance and reliability of behavioural ecotoxicity studies
Behavioural analysis has been attracting significant attention as a broad indicator of sub-lethal toxicity and has secured a place as an important subdiscipline in ecotoxicology. Among the most notable characteristics of behavioural research, compared to other established approaches in sub-lethal ecotoxicology (e.g. reproductive and developmental bioassays), are the wide range of study designs being used and the diversity of endpoints considered. At the same time, environmental hazard and risk assessment, which underpins regulatory decisions to protect the environment from potentially harmful chemicals, often recommends that ecotoxicological data be produced following accepted and validated test guidelines. These guidelines typically do not address behavioural changes, meaning that these, often sensitive, effects are not represented in hazard and risk assessments. Here, we propose a new tool, the EthoCRED evaluation method, for assessing the relevance and reliability of behavioural ecotoxicity data, which considers the unique requirements and challenges encountered in this field. This method and accompanying reporting recommendations are designed to serve as an extension of the "Criteria for Reporting and Evaluating Ecotoxicity Data (CRED)" project. As such, EthoCRED can both accommodate the wide array of experimental design approaches seen in behavioural ecotoxicology, and could be readily implemented into regulatory frameworks as deemed appropriate by policy makers of different jurisdictions to allow better integration of knowledge gained from behavioural testing into environmental protection. Furthermore, through our reporting recommendations, we aim to improve the reporting of behavioural studies in the peer-reviewed literature, and thereby increase their usefulness to inform chemical regulation.
Automatic detection for bioacoustic research: a practical guide from and for biologists and computer scientists
Recent years have seen a dramatic rise in the use of passive acoustic monitoring (PAM) for biological and ecological applications, and a corresponding increase in the volume of data generated. However, data sets are often becoming so sizable that analysing them manually is increasingly burdensome and unrealistic. Fortunately, we have also seen a corresponding rise in computing power and the capability of machine learning algorithms, which offer the possibility of performing some of the analysis required for PAM automatically. Nonetheless, the field of automatic detection of acoustic events is still in its infancy in biology and ecology. In this review, we examine the trends in bioacoustic PAM applications, and their implications for the burgeoning amount of data that needs to be analysed. We explore the different methods of machine learning and other tools for scanning, analysing, and extracting acoustic events automatically from large volumes of recordings. We then provide a step-by-step practical guide for using automatic detection in bioacoustics. One of the biggest challenges for the greater use of automatic detection in bioacoustics is that there is often a gulf in expertise between the biological sciences and the field of machine learning and computer science. Therefore, this review first presents an overview of the requirements for automatic detection in bioacoustics, intended to familiarise those from a computer science background with the needs of the bioacoustics community, followed by an introduction to the key elements of machine learning and artificial intelligence that a biologist needs to understand to incorporate automatic detection into their research. We then provide a practical guide to building an automatic detection pipeline for bioacoustic data, and conclude with a discussion of possible future directions in this field.
The cryptonephridial/rectal complex: an evolutionary adaptation for water and ion conservation
Arthropods have integrated digestive and renal systems, which function to acquire and maintain homeostatically the substances they require for survival. The cryptonephridial complex (CNC) is an evolutionary novelty in which the renal organs and gut have been dramatically reorganised. Parts of the renal or Malpighian tubules (MpTs) form a close association with the surface of the rectum, and are surrounded by a novel tissue, the perinephric membrane, which acts to insulate the system from the haemolymph and thus allows tight regulation of ions and water into and out of the CNC. The CNC can reclaim water and solutes from the rectal contents and recycle these back into the haemolymph. Fluid flow in the MpTs runs counter to flow within the rectum. It is this countercurrent arrangement that underpins its powerful recycling capabilities, and represents one of the most efficient water conservation mechanisms in nature. CNCs appear to have evolved multiple times, and are present in some of the largest and most evolutionarily successful insect groups including the larvae of most Lepidoptera and in a major beetle lineage (Cucujiformia + Bostrichoidea), suggesting that the CNC is an important adaptation. Here we review the knowledge of this remarkable organ system gained over the past 200 years. We first focus on the CNCs of tenebrionid beetles, for which we have an in-depth understanding from physiological, structural and ultrastructural studies (primarily in Tenebrio molitor), which are now being extended by studies in Tribolium castaneum enabled by advances in molecular and microscopy approaches established for this species. These recent studies are beginning to illuminate CNC development, physiology and endocrine control. We then take a broader view of arthropod CNCs, phylogenetically mapping their reported occurrence to assess their distribution and likely evolutionary origins. We explore CNCs from an ecological viewpoint, put forward evidence that CNCs may primarily be adaptations for facing the challenges of larval life, and argue that their loss in many aquatic species could point to a primary function in conserving water in terrestrial species. Finally, by considering the functions of renal and digestive epithelia in insects lacking CNCs, as well as the typical architecture of these organs in relation to one another, we propose that ancestral features of these organs predispose them for the evolution of CNCs.
Trouble on the horizon: anticipating biological invasions through futures thinking
Anticipating future biosecurity threats to prevent their occurrence is the most cost-effective strategy to manage invasive alien species. Yet, biological invasions are complex, highly uncertain processes. High uncertainty drives decision-making away from strategic preventative measures and towards operational outcomes aimed at post-invasion management. The limited success of preventative measures in curbing biological invasions reflects this short-term mindset and decision-makers should instead apply strategic foresight to imagine futures where biosecurity threats are minimised. Here, four major futures thinking tools (environmental scanning, driver-mapping, horizon scanning, and scenario planning) that describe probable, possible, plausible and preferable futures are assessed in terms of their potential to support both research and policy addressing biological invasions. Environmental scanning involves surveying existing data sources to detect signals of emerging alien species through knowledge of changes in either the likelihood or consequences of biological invasions. Several approaches are widely used for biosecurity including automated scans of digital media, consensus-based expert scoring, and prediction markets. Automated systems can be poor at detecting weak signals because of the large volume of 'noise' they generate while expert scoring relies on prior knowledge and so fails to identify unknown unknowns which is also true of prediction markets that work well for quite specific known risks. Driver-mapping uses expert consensus to identify the political, economic, societal, technological, legislative, and environmental forces shaping the future and is a critical component of strategic foresight that has rarely been applied to biological invasions. Considerable potential exists to extend this approach to develop system maps to identify where biosecurity interventions may be most effective and to explore driver complexes to determine megatrends shaping the future of biological invasions. Horizon scanning is a systematic outlook of potential threats and future developments to detect weak signals of emerging issues that exist at the margins of current thinking. Applications have been strongly focused on emerging issues related to research and technological challenges relevant to biosecurity and invasion science. However, most of these emerging issues are already well known in current-day research. Because horizon scanning is based on expert consensus, it needs to embrace a diversity of cultural, gender, and disciplinary diversity more adequately to ensure participants think intuitively and outside of their own subject boundaries. Scenario planning constructs storylines that describe alternative ways the political, economic, social, technological, legislative, and environmental situation might develop in the future. Biological invasion scenario planning has favoured structured approaches such as standardised archetypes and uncertainty matrices, but scope exists to apply more intuitive thinking by using incasting, backcasting, or causal layered analysis. Futures thinking in biological invasions has not engaged with decision-makers or other stakeholders adequately and thus outcomes have been light on policy and management priorities. To date, strategic foresight addressing biological invasions has applied each approach in isolation. Yet, an integrated approach to futures thinking that involves a diverse set of stakeholders in exploring the probable, possible, plausible, and preferable futures relating to biological invasions is crucial to the delivery of strategic biosecurity foresight at both national and global scales.
Protection of the genome and the central exome by peripheral non-coding DNA against DNA damage in health, ageing and age-related diseases
DNA in eukaryotic genomes is under constant assault from both exogenous and endogenous sources, leading to DNA damage, which is considered a major molecular driver of ageing. Fortunately, the genome and the central exome are safeguarded against these attacks by abundant peripheral non-coding DNA. Non-coding DNA codes for small non-coding RNAs that inactivate foreign nucleic acids in the cytoplasm and physically blocks these attacks in the nucleus. Damage to non-coding DNA produced during such blockage is removed in the form of extrachromosomal circular DNA (eccDNA) through nucleic pore complexes. Consequently, non-coding DNA serves as a line of defence for the exome against DNA damage. The total amount of non-coding DNA/heterochromatin declines with age, resulting in a decrease in both physical blockage and eccDNA exclusion, and thus an increase in the accumulation of DNA damage in the nucleus during ageing and in age-related diseases. Here, we summarize recent evidence supporting a protective role of non-coding DNA in healthy and pathological states and argue that DNA damage is the proximate cause of ageing and age-related genetic diseases. Strategies aimed at strengthening the protective role of non-coding DNA/heterochromatin could potentially offer better systematic protection for the dynamic genome and the exome against diverse assaults, reduce the burden of DNA damage to the exome, and thus slow ageing, counteract age-related genetic diseases and promote a healthier life for individuals.