From success to persistence: Identifying an evolutionary regime shift in the diverse Paleozoic aquatic arthropod group Eurypterida,driven by the Devonian biotic crisis

Mass extinctions have altered the trajectory of evolution a number of times over the Phanerozoic. During these periods of biotic upheaval a different selective regime appears to operate, although it is still unclear whether consistent survivorship rules apply across different extinction events. We compare variations in diversity and disparity across the evolutionary history of a major Paleozoic arthropod group, the Eurypterida. Using these data, we explore the group's transition from a successful, dynamic clade to a stagnant persistent lineage, pinpointing the Devonian as the period during which this evolutionary regime shift occurred. The late Devonian biotic crisis is potentially unique among the “Big Five” mass extinctions in exhibiting a drop in speciation rates rather than an increase in extinction. Our study reveals eurypterids show depressed speciation rates throughout the Devonian but no abnormal peaks in extinction. Loss of morphospace occupation is random across all Paleozoic extinction events; however, differential origination during the Devonian results in a migration and subsequent stagnation of occupied morphospace. This shift appears linked to an ecological transition from euryhaline taxa to freshwater species with low morphological diversity alongside a decrease in endemism. These results demonstrate the importance of the Devonian biotic crisis in reshaping Paleozoic ecosystems.     

Lazarus ecology: Recovering the distribution and migratory patterns of the extinct Carolina parakeet

The study of the ecology and natural history of species has traditionally ceased when a species goes extinct, despite the benefit to current and future generations of potential findings. We used the extinct Carolina parakeet as a case study to develop a framework investigating the distributional limits, subspecific variation, and migratory habits of this species as a means to recover important information about recently extinct species. We united historical accounts with museum collections to develop an exhaustive, comprehensive database of every known occurrence of this once iconic species. With these data, we combined species distribution models and ordinal niche comparisons to confront multiple conjectured hypotheses about the parakeet's ecology with empirical data on where and when this species occurred. Our results demonstrate that the Carolina parakeet's range was likely much smaller than previously believed, that the eastern and western subspecies occupied different climatic niches with broad geographical separation, and that the western subspecies was likely a seasonal migrant while the eastern subspecies was not. This study highlights the novelty and importance of collecting occurrence data from published observations on extinct species, providing a starting point for future investigations of the factors that drove the Carolina parakeet to extinction. Moreover, the recovery of lost autecological knowledge could benefit the conservation of other parrot species currently in decline and would be crucial to the success of potential de‐extinction efforts for the Carolina parakeet.     

More losers than winners: investigating Anthropocene defaunation through the diversity of population trends

The global-scale decline of animal biodiversity (‘defaunation’) represents one of the most alarming consequences of human impacts on the planet. The quantification of this extinction crisis has traditionally relied on the use of IUCN Red List conservation categories assigned to each assessed species. This approach reveals that a quarter of the world's animal species are currently threatened with extinction, and ~1% have been declared extinct. However, extinctions are preceded by progressive population declines through time that leave demographic ‘footprints’ that can alert us about the trajectories of species towards extinction. Therefore, an exclusive focus on IUCN conservation categories, without consideration of dynamic population trends, may underestimate the true extent of the processes of ongoing extinctions across nature. In fact, emerging evidence (e.g. the Living Planet Report), reveals a widespread tendency for sustained demographic declines (an average 69% decline in population abundances) of species globally. Yet, animal species are not only declining. Many species worldwide exhibit stable populations, while others are even thriving. Here, using population trend data for >71,000 animal species spanning all five groups of vertebrates (mammals, birds, reptiles, amphibians and fishes) and insects, we provide a comprehensive global-scale assessment of the diversity of population trends across species undergoing not only declines, but also population stability and increases. We show a widespread global erosion of species, with 48% undergoing declines, while 49% and 3% of species currently remain stable or are increasing, respectively. Geographically, we reveal an intriguing pattern similar to that of threatened species, whereby declines tend to concentrate around tropical regions, whereas stability and increases show a tendency to expand towards temperate climates. Importantly, we find that for species currently classed by the IUCN Red List as ‘non-threatened’, 33% are declining. Critically, in contrast with previous mass extinction events, our assessment shows that the Anthropocene extinction crisis is undergoing a rapid biodiversity imbalance, with levels of declines (a symptom of extinction) greatly exceeding levels of increases (a symptom of ecological expansion and potentially of evolution) for all groups. Our study contributes a further signal indicating that global biodiversity is entering a mass extinction, with ecosystem heterogeneity and functioning, biodiversity persistence, and human well-being under increasing threat.     

Evaluating the predicted extinction risk of living amphibian species with the fossil record

Bridging the gap between the fossil record and conservation biology has recently become of great interest. The enormous number of documented extinctions across different taxa can provide insights into the extinction risk of living species. However, few studies have explored this connection. We used generalised boosted modelling to analyse the impact of several traits that are assumed to influence extinction risk on the stratigraphic duration of amphibian species in the fossil record. We used this fossil‐calibrated model to predict the extinction risk for living species. We observed a high consensus between our predicted species durations and the current IUCN Red List status of living amphibian species. We also found that today's Data Deficient species are mainly predicted to experience short durations, hinting at their likely high threat status. Our study suggests that the fossil record can be a suitable tool for the evaluation of current taxa‐specific Red Listing status.     

Darwin's error? Patrick Matthew and the catastrophic nature of the geologic record

In 1831, the Scottish horticulturalist Patrick Matthew (1790–1874) published a clear statement of the law of natural selection in an Appendix to his book Naval Timber and Arboriculture, which both Darwin and Wallace later acknowledged. Matthew, however, was a catastrophist, and he presented natural selection within the contemporary view that relatively long intervals of environmental stability were episodically punctuated by catastrophic mass extinctions of life. Modern studies support a similar picture of the division of geologic time into long periods of relative evolutionary stability ended by sudden extinction events. Mass extinctions are followed by recovery intervals during which surviving taxa radiate into vacated niches. This modern punctuated view of evolution and speciation is much more in line with Matthew's episodic catastrophism than the classical Lyellian–Darwinian gradualist view.     

Body length of bony fishes was not a selective factor during the biggest mass extinction of all time

The Permo‐Triassic mass extinction devastated life on land and in the sea, but it is not clear why some species survived and others went extinct. One explanation is that lineage loss during mass extinctions is a random process in which luck determines which species survive. Alternatively, a phylogenetic signal in extinction may indicate a selection process operating on phenotypic traits. Large body size has often emerged as an extinction risk factor in studies of modern extinction risk, but this is not so commonly the case for mass extinctions in deep time. Here, we explore the evolution of non‐teleostean Actinopterygii (bony fishes) from the Devonian to the present day, and we concentrate on the Permo‐Triassic mass extinction. We apply a variety of time‐scaling metrics to date the phylogeny, and show that diversity peaked in the latest Permian and declined severely during the Early Triassic. In line with previous evidence, we find the phylogenetic signal of extinction increases across the mass extinction boundary: extinction of species in the earliest Triassic is more clustered across phylogeny compared to the more randomly distributed extinction signal in the late Permian. However, body length plays no role in differential survival or extinction of taxa across the boundary. In the case of fishes, size did not determine which species survived and which went extinct, but phylogenetic signal indicates that the mass extinction was not a random field of bullets.     

Reproductive failure: a new paradigm for extinction

Extinction was recognized as a scientific fact 200 years ago, although no adequate paradigm has emerged to explain the process. Prevailing theory has focused on ‘cause(s)’ of extinction but has neglected ‘effect’ and ‘mechanism’. These omissions preclude the formulation of a functional paradigm necessary for remedial action in response to the impending anthropogenic mediated, worldwide extinction crisis. The new paradigm is defined as the multi‐generational, attritional loss of reproductive fitness. Stabilizing selection continuously adapts species to specific ecosystems, which often results in highly evolved species prone to extinction when environments shift. Some species survive by tracking the declining palaeoclimates in which they presumably evolved, often becoming relicts prior to extinction. Compelling new evidence shows that even mass extinctions are largely a result of environmental change leading to widespread, attritional reproductive decline, rather than a result of instantaneous global catastrophes.     

Body size evolution in an old insect order: No evidence for Cope's Rule in spite of fitness benefits of large size

We integrate field data and phylogenetic comparative analyses to investigate causes of body size evolution and stasis in an old insect order: odonates (“dragonflies and damselflies”). Fossil evidence for “Cope's Rule” in odonates is weak or nonexistent since the last major extinction event 65 million years ago, yet selection studies show consistent positive selection for increased body size among adults. In particular, we find that large males in natural populations of the banded demoiselle (Calopteryx splendens) over several generations have consistent fitness benefits both in terms of survival and mating success. Additionally, there was no evidence for stabilizing or conflicting selection between fitness components within the adult life‐stage. This lack of stabilizing selection during the adult life‐stage was independently supported by a literature survey on different male and female fitness components from several odonate species. We did detect several significant body size shifts among extant taxa using comparative methods and a large new molecular phylogeny for odonates. We suggest that the lack of Cope's rule in odonates results from conflicting selection between fitness advantages of large adult size and costs of long larval development. We also discuss competing explanations for body size stasis in this insect group.     

Extinction

A significant proportion of conservationists' work is directed towards efforts to save disappearing species. This relies upon the belief that species extinction is undesirable. When justifications are offered for this belief, they very often rest upon the assumption that extinction brought about by humans is different in kind from other forms of extinction. This paper examines this assumption and reveals that there is indeed good reason to suppose current anthropogenic extinctions to be different in kind from extinctions brought about at other times or by other factors. Having considered – and rejected – quantity and rate of extinction as useful distinguishing factors, four alternative arguments are offered, each identifying a way in which anthropogenic extinction is significantly different from other forms of extinction, even mass extinction: (1) Humans are a different kind of natural cause from other causes of extinction; (2) Extinctions brought about by humans are uniquely persistent; (3) Anthropogenic extinctions are effectively random whereas past mass extinctions are rule-bound; (4) The impact of the current anthropogenic extinction event differs from the impact of other extinction events of the past, such that future recovery may not follow past patterns. Together, these four arguments suggest that the present-day extinction event brought about by humans may be unprecedented and that we cannot clearly extrapolate from past to present recovery from extinctions. Although insufficient as justification for the claim that present-day extinctions are undesirable, the arguments provide some ammunition for conservationists' conviction that species extinction – in which humans play an accelerating role – ought to be prevented.     

The genetic architecture of ecological speciation and the association with signatures of selection in natural lake whitefish (Coregonus sp. Salmonidae) species pairs

Adaptive evolutionary change is contingent on variation and selection; thus, understanding adaptive divergence and ultimately speciation requires information on both the genetic basis of adaptive traits as well as an understanding of the role of divergent natural selection on those traits. The lake whitefish (Coregonus clupeaformis) consists of several sympatric "dwarf" (limnetic) and normal (benthic) species pairs that co-inhabit northern postglacial lakes. These young species pairs have evolved independently and display parallelism in life history, behavioral, and morphological divergence associated with the use of distinct trophic resources. We identified phenotype-environment associations and determined the genetic architecture and the role of selection modulating population genetic divergence in sympatric dwarf and normal lake whitefish. The genetic architecture of 9 adaptive traits was analyzed in 2 hybrid backcrosses individually phenotyped throughout their life history. Significant quantitative trait loci (QTL) were associated with swimming behavior (habitat selection and predator avoidance), growth rate, morphology (condition factor and gill rakers), and life history (onset of maturity and fecundity). Genome scans among 4 natural sympatric pairs, using loci segregating in the map, revealed a signature of selection for 24 loci. Loci exhibiting a signature of selection were associated with QTL relative to other regions of the genome more often than expected by chance alone. Two parallel QTL outliers for growth and condition factor exhibited segregation distortion in both mapping families, supporting the hypothesis that adaptive divergence contributing to parallel reductions of gene flow among natural populations may cause genetic incompatibilities. Overall, these findings offer evidence that the genetic architecture of ecological speciation is associated with signatures of selection in nature, providing strong support for the hypothesis that divergent natural selection is currently maintaining adaptive differentiation and promoting ecological speciation in lake whitefish species pairs.     

Why Darwin would have loved evolutionary game theory

Humans have marvelled at the fit of form and function, the way organisms'' traits seem remarkably suited to their lifestyles and ecologies. While natural selection provides the scientific basis for the fit of form and function, Darwin found certain adaptations vexing or particularly intriguing: sex ratios, sexual selection and altruism. The logic behind these adaptations resides in frequency-dependent selection where the value of a given heritable phenotype (i.e. strategy) to an individual depends upon the strategies of others. Game theory is a branch of mathematics that is uniquely suited to solving such puzzles. While game theoretic thinking enters into Darwin''s arguments and those of evolutionists through much of the twentieth century, the tools of evolutionary game theory were not available to Darwin or most evolutionists until the 1970s, and its full scope has only unfolded in the last three decades. As a consequence, game theory is applied and appreciated rather spottily. Game theory not only applies to matrix games and social games, it also applies to speciation, macroevolution and perhaps even to cancer. I assert that life and natural selection are a game, and that game theory is the appropriate logic for framing and understanding adaptations. Its scope can include behaviours within species, state-dependent strategies (such as male, female and so much more), speciation and coevolution, and expands beyond microevolution to macroevolution. Game theory clarifies aspects of ecological and evolutionary stability in ways useful to understanding eco-evolutionary dynamics, niche construction and ecosystem engineering. In short, I would like to think that Darwin would have found game theory uniquely useful for his theory of natural selection. Let us see why this is so.     

Confronting a biome crisis: global disparities of habitat loss and protection

Human impacts on the natural environment have reached such proportions that in addition to an ‘extinction crisis’, we now also face a broader ‘biome crisis’. Here we identify the world's terrestrial biomes and, at a finer spatial scale, ecoregions in which biodiversity and ecological function are at greatest risk because of extensive habitat conversion and limited habitat protection. Habitat conversion exceeds habitat protection by a ratio of 8 : 1 in temperate grasslands and Mediterranean biomes, and 10 : 1 in more than 140 ecoregions. These regions include some of the most biologically distinctive, species rich ecosystems on Earth, as well as the last home of many threatened and endangered species. Confronting the biome crisis requires a concerted and comprehensive response aimed at protecting not only species, but the variety of landscapes, ecological interactions, and evolutionary pressures that sustain biodiversity, generate ecosystem services, and evolve new species in the future.     

盖娅假说:在争议中发展

自从1972年Lovelock提出盖娅假说已经过去了40年,但围绕它的争议却从未停止过。盖娅假说在反对者的批评中与支持者的证明中不断发展。当前,最极端形式的盖娅假说基本上已被摒弃,尤其是那种明显带有目的论的说法。弱盖娅提出的"有机体可以影响他们的环境,有机体与环境的反馈耦合可以塑造两者的进化"这两个观点也已经是普遍接受的事实。除此之外,盖娅假说提出的其他3个命题却饱受争议。(1)内在平衡的盖娅:生物调节反馈有助于环境的内在平衡。反对者认为,生物反馈稳定全球环境的说法,与冰芯记录和大量的气候反馈研究结果相矛盾的。支持者认为,地球生物-环境系统的内在平衡可以产生于正负反馈的混合。盖娅假说关心的是地球几十亿年的历史,盖娅假说在较短时间尺度内可证伪,并不意味着其在较长时间尺度内也可证伪。(2)最优的盖娅:生物调节环境,使环境更加适合生物的生存。关于有机体的繁荣主要是由于他们对环境的改变,还是由于他们对环境的适应,目前尚未有结论。但盖娅的支持者认为,当生物-环境系统受到干扰或崩溃时,主导过程将显现。拥有较强环境反馈的系统,将易于快速过渡到新的状态,而由适应主导的过程将改变得较为平缓。反对者同意生物通过生物调节作用影响环境条件以使自身受益,但是生物首先要适应环境条件通过自然选择才能得以繁荣发展的。地球形成这样的环境条件,很可能纯粹是一种运气。(3)自然选择的盖娅:生物调节反馈产生于达尔文式的自然选择。反对者认为,"自然选择支持促进生命效应"的说法并非普遍有效,只有当遗传特征赋予携带者繁殖优势时,自然选择才会支持它。自然选择是机制,而非原则。支持者认为自然选择并不是盖娅系统环境调节的必要条件;基于副产品的自然选择,可以解决许多进化论学者提出的物种合作中的欺骗问题;自然选择并不总是支持促进生命的效应,但在当遗传特征使携带者相对非携带者受益时,自然选择可以使特征携带者产生进化优势。虽然争议依然存在并将持续下去,但作为假说生产者,盖娅假说已经证明了它的价值。但是在人类活动对生物圈影响不断增强的背景下,盖娅假说必须与人类活动相结合,否则必然走向衰落,并被其他理论或假说所替代。在此基础上,未来盖娅假说的研究者们需要继续努力探索可以应用于生物圈的一般性原则,并坚持系统性的思考方法。在具体的方法方面,可以利用系统度量指标;建立新的模型,尤其是建立关于生物地球化学循环过程的机理模型;搞清楚不同尺度过程的成本与收益。     

Body size changes in bivalves of the family Limidae in the aftermath of the end‐Triassic mass extinction: the Brobdingnag effect

Reduction in body size of organisms following mass extinctions is well‐known and often ascribed to the Lilliput effect. This phenomenon is expressed as a temporary body size reduction within surviving species. Despite its wide usage the term is often loosely applied to any small post‐extinction taxa. Here we assess the size of bivalves of the family Limidae (Rafineque) prior to, and in the aftermath of, the end‐Triassic mass extinction event. Of the species studied only one occurs prior to the extinction event, though is too scarce to test for the Lilliput effect. Instead, newly evolved species originate at small body sizes and undergo a within‐species size increase, most dramatically demonstrated by Plagiostoma giganteum (Sowerby) which, over two million years, increases in size by 179%. This trend is seen in both field and museum collections. We term this within‐species size increase of newly originated species in the aftermath of mass extinction, the Brobdingnag effect, after the giants that were contemporary with the Lilliputians in Swift's Gulliver's Travels. The size increase results from greater longevity and faster growth rates. The cause of the effect is unclear, although it probably relates to improved environmental conditions. Oxygen‐poor conditions in the Early Jurassic are associated with populations of smaller body size caused by elevated juvenile mortality but these are local/regional effects that do not alter the long‐term, size increase. Although temperature‐size relationships exist for many organisms (Temperature‐Size Rule and Bergmann's Rule), the importance of this is unclear here because of a poorly known Early Jurassic temperature record.     

Mammal extinctions and the increasing isolation of humans on the tree of life

A sixth great mass extinction is ongoing due to the direct and indirect effects of human pressures. However, not all lineages are affected equally. From an anthropocentric perspective, it is often purported that humans hold a unique place on Earth. Here, we show that our current impacts on the natural world risk realizing that expectation. We simulated species loss on the mammalian phylogenetic tree, informed by species current extinction risks. We explored how Homo sapiens could become isolated in the tree if species currently threatened with extinction disappeared. We analyzed correlates of mammal extinctions risks that may drive this isolation pattern. We show that, within mammals, and more particularly within primates, extinction risks increase with the number of known threat types, and decrease with geographic range size. Extinctions increase with species body mass, trophic level, and the median longitudinal extent of each species range in mammals but not within primates. The risks of extinction are frequently high among H. sapiens close relatives. Pruning threatened primates, including apes (Hominidae, Hylobatidae), from the tree of life will lead to our species being among those with the fewest close relatives. If no action is taken, we will thus not only lose crucial biodiversity for the preservation of Earth ecosystems, but also a key living reference to what makes us human.     

Sexual selection predicts the persistence of populations within altered environments

The effect of sexual selection on species persistence remains unclear. The cost of bearing ornaments or armaments might increase extinction risk, but sexual selection can also enhance the spread of beneficial alleles and increase the removal of deleterious alleles, potentially reducing extinction risk. Here we investigate the effect of sexual selection on species persistence in a community of 34 species of dung beetles across a gradient of environmental disturbance ranging from old growth forest to oil palm plantation. Horns are sexually selected traits used in contests between males, and we find that both horn presence and relative size are strongly positively associated with species persistence and abundance in altered habitats. Testes mass, an indicator of post‐copulatory selection, is, however, negatively linked with the abundance of species within the most disturbed habitats. This study represents the first evidence from a field system of a population‐level benefit from pre‐copulatory sexual selection.     

Evolutionary responses of natives to introduced species: what do introductions tell us about natural communities?

Biological invasions dramatically affect the distribution, abundance and reproduction of many native species. Because of these ecological effects, exotic species can also influence the evolution of natives exposed to novel interactions with invaders. Evolutionary changes in natives in response to selection from exotics are usually overlooked, yet common responses include altered anti-predator defenses, changes in the spectrum of resources and habitats used, and other adaptations that allow native populations to persist in invaded areas. Whether a native population is capable of responding evolutionarily to selection from invaders will depend on the demographic impact of the invader, the genetic architecture and genetic variability of the native population and potentially the history of previous invasions. In some cases, natives will fail to evolve or otherwise adapt, and local or global extinction will result. In other cases, adaptive change in natives may diminish impacts of invaders and potentially promote coexistence between invaders and natives. Here, we review the evidence for evolutionary responses of native species to novel community members. We also discuss how the effects of introduced species may differ from those caused by natural range expansions of native species. Notably, introduced species may come from remote biotas with no previous evolutionary history with the native community. In addition, the rate of addition of introduced species into communities is much greater than all but the most extreme cases of historical biotic exchange. Understanding the evolutionary component of exotic/native species interactions is critical to recognizing the long-term impacts of biological invasions, and to understanding the role of evolutionary processes in the assembly and dynamics of natural communities.     

Archosauromorph extinction selectivity during the Triassic–Jurassic mass extinction

Many traits have been linked to extinction risk among modern vertebrates, including mode of life and body size. However, previous work has indicated there is little evidence that body size, or any other trait, was selective during past mass extinctions. Here, we investigate the impact of the Triassic–Jurassic mass extinction on early Archosauromorpha (basal dinosaurs, crocodylomorphs and their relatives) by focusing on body size and other life history traits. We built several new archosauromorph maximum‐likelihood supertrees, incorporating uncertainty in phylogenetic relationships. These supertrees were then employed as a framework to test whether extinction had a phylogenetic signal during the Triassic–Jurassic mass extinction, and whether species with certain traits were more or less likely to go extinct. We find evidence for phylogenetic signal in extinction, in that taxa were more likely to become extinct if a close relative also did. However, there is no correlation between extinction and body size, or any other tested trait. These conclusions add to previous findings that body size, and other traits, were not subject to selection during mass extinctions in closely‐related clades, although the phylogenetic signal in extinction indicates that selection may have acted on traits not investigated here.     

The natural selection of metabolism explains curvature in allometric scaling

I simulate the natural selection of metabolism and mass to explain the curvature in the metabolic allometry for placental and marsupial mammals. The simulation model starts with a single ancestor in each clade at the Cretaceous–Palaeogene boundary 65 million years ago. The release of inter‐specific competition by the extinction of dinosaurs make it possible for each clade to diversify into a multitude of species across a wide range of empty niches. The selection of mass in these species depends on the net assimilated energy that depends on 1) the handling of the resources in the different niches, and on 2) mass‐specific metabolism that defines the pace of the handling process. The model is fitted to explain the maximum observed body masses over time and the current inter‐specific allometry for metabolism. The selection of mass‐specific metabolism is found to bend the metabolic allometry over time, even when all species have the same selection on the per‐generation time‐scale of natural selection. This is because the smaller species evolve over a larger number of generations than the larger species. The strongest curvature is in the placental clade, where the estimated rate of exponential increase in mass‐specific metabolism is 9.3 × 10^?9 (95% CI: 7.3 × 10^?9 – 1.1 × 10^?8) on the per‐generation time‐scale. This is an order of magnitude larger than the estimate for marsupials, in agreement with an average metabolism that is 30% larger in placentals relative to marsupials of similar size.     

Host–parasite coevolution: Partitioning the effects of natural selection and environmental change using coupled Price equations

George Price showed how the effects of natural selection and environmental change could be mathematically partitioned. This partitioning may be especially useful for understanding host–parasite coevolution, where each species represents the environment for the other species. Here, we use coupled Price equations to study this kind of antagonistic coevolution. We made the common assumption that parasites must genetically match their host''s genotype to avoid detection by the host''s self/nonself recognition system, but we allowed for the possibility that non‐matching parasites have some fitness. Our results show how natural selection on one species results in environmental change for the other species. Numerical iterations of the model show that these environmental changes can periodically exceed the changes in mean fitness due to natural selection, as suggested by R.A. Fisher. Taken together, the results give an algebraic dissection of the eco‐evolutionary feedbacks created during host–parasite coevolution.