Predicting loss of evolutionary history: Where are we?

The Earth's evolutionary history is threatened by species loss in the current sixth mass extinction event in Earth's history. Such extinction events not only eliminate species but also their unique evolutionary histories. Here we review the expected loss of Earth's evolutionary history quantified by phylogenetic diversity (PD) and evolutionary distinctiveness (ED) at risk. Due to the general paucity of data, global evolutionary history losses have been predicted for only a few groups, such as mammals, birds, amphibians, plants, corals and fishes. Among these groups, there is now empirical support that extinction threats are clustered on the phylogeny; however this is not always a sufficient condition to cause higher loss of phylogenetic diversity in comparison to a scenario of random extinctions. Extinctions of the most evolutionarily distinct species and the shape of phylogenetic trees are additional factors that can elevate losses of evolutionary history. Consequently, impacts of species extinctions differ among groups and regions, and even if global losses are low within large groups, losses can be high among subgroups or within some regions. Further, we show that PD and ED are poorly protected by current conservation practices. While evolutionary history can be indirectly protected by current conservation schemes, optimizing its preservation requires integrating phylogenetic indices with those that capture rarity and extinction risk. Measures based on PD and ED could bring solutions to conservation issues, however they are still rarely used in practice, probably because the reasons to protect evolutionary history are not clear for practitioners or due to a lack of data. However, important advances have been made in the availability of phylogenetic trees and methods for their construction, as well as assessments of extinction risk. Some challenges remain, and looking forward, research should prioritize the assessment of expected PD and ED loss for more taxonomic groups and test the assumption that preserving ED and PD also protects rare species and ecosystem services. Such research will be useful to inform and guide the conservation of Earth's biodiversity and the services it provides.     

The future of evolutionary diversity in reef corals

One-third of the world''s reef-building corals are facing heightened extinction risk from climate change and other anthropogenic impacts. Previous studies have shown that such threats are not distributed randomly across the coral tree of life, and future extinctions have the potential to disproportionately reduce the phylogenetic diversity of this group on a global scale. However, the impact of such losses on a regional scale remains poorly known. In this study, we use phylogenetic metrics in conjunction with geographical distributions of living reef coral species to model how extinctions are likely to affect evolutionary diversity across different ecoregions. Based on two measures—phylogenetic diversity and phylogenetic species variability—we highlight regions with the largest losses of evolutionary diversity and hence of potential conservation interest. Notably, the projected loss of evolutionary diversity is relatively low in the most species-rich areas such as the Coral Triangle, while many regions with fewer species stand to lose much larger shares of their diversity. We also suggest that for complex ecosystems like coral reefs it is important to consider changes in phylogenetic species variability; areas with disproportionate declines in this measure should be of concern even if phylogenetic diversity is not as impacted. These findings underscore the importance of integrating evolutionary history into conservation planning for safeguarding the future diversity of coral reefs.     

Threatened reef corals of the world

A substantial proportion of the world's living species, including one-third of the reef-building corals, are threatened with extinction and in pressing need of conservation action. In order to reduce biodiversity loss, it is important to consider species' contribution to evolutionary diversity along with their risk of extinction for the purpose of setting conservation priorities. Here I reconstruct the most comprehensive tree of life for the order Scleractinia (1,293 species) that includes all 837 living reef species, and employ a composite measure of phylogenetic distinctiveness and extinction risk to identify the most endangered lineages that would not be given top priority on the basis of risk alone. The preservation of these lineages, not just the threatened species, is vital for safeguarding evolutionary diversity. Tests for phylogeny-associated patterns show that corals facing elevated extinction risk are not clustered on the tree, but species that are susceptible, resistant or resilient to impacts such as bleaching and disease tend to be close relatives. Intensification of these threats or extirpation of the endangered lineages could therefore result in disproportionate pruning of the coral tree of life.     

Conservation of reef corals in the South China Sea based on species and evolutionary diversity

The South China Sea in the Central Indo-Pacific is a large semi-enclosed marine region that supports an extraordinary diversity of coral reef organisms (including stony corals), which varies spatially across the region. While one-third of the world’s reef corals are known to face heightened extinction risk from global climate and local impacts, prospects for the coral fauna in the South China Sea region amidst these threats remain poorly understood. In this study, we analyse coral species richness, rarity, and phylogenetic diversity among 16 reef areas in the region to estimate changes in species and evolutionary diversity during projected anthropogenic extinctions. Our results show that richness, rarity, and phylogenetic diversity differ considerably among reef areas in the region, and that their outcomes following projected extinctions cannot be predicted by species diversity alone. Although relative rarity and threat levels are high in species-rich areas such as West Malaysia and the Philippines, areas with fewer species such as northern Vietnam and Paracel Islands stand to lose disproportionately large amounts of phylogenetic diversity. Our study quantifies various biodiversity components of each reef area to inform conservation planners and better direct sparse resources to areas where they are needed most. It also provides a critical biological foundation for targeting reefs that should be included in a regional network of marine protected areas in the South China Sea.     

Reef coral diversity and global change

Regional anthropogenic processes such as pollution, dredging, and overfishing on coral reefs currently threaten the biodiversity of stony corals and other reef-associated organisms. Global climate change may interact with anthropogenic processes to create additional impacts on coral diversity in the near future. In order to predict these changes, it is necessary to understand the magnitude and causes of variation in scleractinian coral diversity throughout their 240 million year history. The fossil record documents long periods of speciation in corals, interrupted repeatedly by events of mass extinction. Some of these events relate clearly to changes in global climate. Diversity in reef corals has increased since their last period of extinction at the end of the Cretaceous (65 My bp ), and is still rising. During the last 8 million years, the fragmentation of the once pantropical Tethys Sea separated corals into two major biogeographical provinces. Periods of glaciation also have caused major changes in sea level and temperature. Accumulated evidence supports the theory that relative habitat area and changing patterns of oceanic circulation are mainly responsible for the two observed centres of recent coral diversity at the western tropical margins of the Atlantic and Pacific oceans. At predicted rates of climate change in the near future, coral reefs are likely to survive as an ecosystem. Increases in sea level may actually benefit corals and lead to regional increases in diversity if new habitat area on back reefs is opened to increased water circulation and thus coral dispersal. Rising temperature may cause higher rates of coral mortality and even local extinction in isolated, small populations such as those on oceanic islands. The effects of increases in ultraviolet radiation (UV) are largely unknown, but likely to be negative. UV may damage planktonic coral propagules in oceanic surface waters and thus decrease rates of gene flow between coral populations. This may result in increased local extinctions, again with the strongest impact on widely separated reefs with small coral populations. The largest threats to coral diversity are regional anthropogenic impacts, which may interact with global climate change to exacerbate rates of local species extinctions. Centres of high reef coral diversity coincide with human population centres in south-east Asia and the Caribbean, and thus the greatest potential for species loss lies in these geographical areas.     

The history of Devonian-Carboniferous reef communities: Extinctions,effects, recovery

Summary Analysis of the taxonomic composition, diversity and guild structure of five “typical” reef and mud mound communities ranging in age from Late Devonian-Early Carboniferous indicates that each of these aspects of community organization changed dramatically in relation to three extinction events. These events include a major or mass extinction at the end of the Frasnian; reef communities were also effected by less drastic end-Givetian and mid-late Famennian extinctions of reef-building higher taxa. Peak Paleozoic generic diversities for reef-building stromatoporoids and rugose corals occurred in the Eifelian-Givetian; reef-building calcareous algal taxa were longranging with peak diversity in the Devonian. These three higher taxa dominated all reef-building guilds (Constructor, Binder, Baffler) in the Frasnian and formed fossil reef communities with balanced guild structures. The extinction of nearly all reef-building stromatoporoids and rugose corals at the end of the Frasnian and the survival of nearly all calcareous algac produced mid-late Famennian reef communities dominated by the Binder Guild. Despite the survival of most calcareous algae and tabulate corals, the mid-late Famennian extinction of all remaining Paleozoic stromatoporoids and nearly all shelf-dwelling Rugosa brought the already diminished Devonian reef-building to a halt. These Devonian extinctions differ from mass extinctions by the absence of a statistically significant drop in taxonomic diversity and by their successional and cumulative effects on reef communities. Tournaisian mud mounds contain communities markedly different from the frame-building communities in Late Devonian and Visean reefs. Mound-building biotas consist of an unusual association dominated by erect, weakly skeletonized members of the Baffler Guild (chiefly fenestrate Bryozoa; Pelmatozoa) and laterally expanded, mud-binding algae/stromatolites and reptant Bryozoa. The initial recovery to reefs with skeletal frameworks in the Visean was largely due to the re-appearance of new species of abundant colonial rugose corals (Constructor Guild) and fenestrate Bryozoa. This Frasnian-Visean evolution in the taxonomic composition and structure of the reef-building guilds is also expressed by abrupt changes in biofacies and petrology of the reef limestones they produced. Thus, “typical” Frasnian reef limestones with balanced guild structures are framestones-boundstones-bafflestones, Famennian reefs are predominantly boundstones, Tournaisian mud mounds are bafflestones and Visean reefs are bafflestones-framestones.     

Impact of the Late Triassic mass extinction on functional diversity and composition of marine ecosystems

Mass extinctions have profoundly influenced the history of life, not only through the death of species but also through changes in ecosystem function and structure. Importantly, these events allow us the opportunity to study ecological dynamics under levels of environmental stress for which there are no recent analogues. Here, we examine the impact and selectivity of the Late Triassic mass extinction event on the functional diversity and functional composition of the global marine ecosystem, and test whether post‐extinction communities in the Early Jurassic represent a regime shift away from pre‐extinction communities in the Late Triassic. Our analyses show that, despite severe taxonomic losses, there is no unequivocal loss of global functional diversity associated with the extinction. Even though no functional groups were lost, the extinction event was, however, highly selective against some modes of life, in particular sessile suspension feeders. Although taxa with heavily calcified skeletons suffered higher extinction than other taxa, lightly calcified taxa also appear to have been selected against. The extinction appears to have invigorated the already ongoing faunal turnover associated with the Mesozoic Marine Revolution. The ecological effects of the Late Triassic mass extinction were preferentially felt in the tropical latitudes, especially amongst reefs, and it took until the Middle Jurassic for reef ecosystems to fully recover to pre‐extinction levels.     

On the potential for ocean acidification to be a general cause of ancient reef crises

Anthropogenic rise in the carbon dioxide concentration in the atmosphere leads to global warming and acidification of the oceans. Ocean acidification (OA) is harmful to many organisms but especially to those that build massive skeletons of calcium carbonate, such as reef corals. Here, we test the recent suggestion that OA leads not only to declining calcification of reef corals and reduced growth rates of reefs but may also have been a trigger of ancient reef crises and mass extinctions in the sea. We analyse the fossil record of biogenic reefs and marine organisms to (1) assess the timing and intensity of ancient reef crises, (2) check which reef crises were concurrent with inferred pulses of carbon dioxide concentrations and (3) evaluate the correlation between reef crises and mass extinctions and their selectivity in terms of inferred physiological buffering. We conclude that four of five global metazoan reef crises in the last 500 Myr were probably at least partially governed by OA and rapid global warming. However, only two of the big five mass extinctions show geological evidence of OA.     

Anthropogenic stressors and riverine fish extinctions

Human activities are often implicated in the contemporary extinction of contemporary species. Concerning riverine fishes, the major biotic and abiotic threats widely cited include introduction of non-native species, habitat fragmentation and homogenization in stream flow dynamics due to the damming of rivers, dumping of organic loadings, degradation of the riverine habitat by agricultural practices and water abstraction for human and agricultural consumption. However, few studies have evaluated the role of each of these threats on fish extinction at large spatial scales. Focusing on Western Europe and the USA, two of the most heavily impacted regions on Earth, we quantify fish species loss per river basin and evaluate for the first time to what extent, if any, these threats have been promoting fish extinctions. We show that mean fish extinction rates during the last 110 years in both continents is ∼112 times higher than calculated natural extinction rates. However, we identified only weak effects of our selected anthropogenic stressors on fish extinctions. Only river fragmentation by dams and percentage of non-native species seem to be significant, although weak, drivers of fish species extinction. In our opinion, the most probable explanation for the weak effects found here comes from limitations of both biological and threats datasets currently available. Obtaining realistic estimates on both extinctions and anthropogenic threats in individual river basins is thus urgently needed.     

Phylogenetically clustered extinction risks do not substantially prune the Tree of Life

Anthropogenic activities have increased the rate of biological extinction many-fold. Recent empirical studies suggest that projected extinction may lead to extensive loss to the Tree of Life, much more than if extinction were random. One suggested cause is that extinction risk is heritable (phylogenetically patterned), such that entire higher groups will be lost. We show here with simulation that phylogenetically clustered extinction risks are necessary but not sufficient for the extensive loss of phylogenetic diversity (PD) compared to random extinction. We simulated Yule trees and evolved extinction risks at various levels of heritability (measured using Pagel's λ). At most levels of heritability (λ in range of 0 to 10), mean values of extinction risk (range 0.25 to 0.75), tree sizes (64 to 128 tips), tree balance and temporal heterogeneity of diversification rates (Yule and coalescent trees), extinction risks do not substantially increase the loss of PD in these trees when compared to random extinction. The maximum loss of PD (20% above random) was only associated with the combination of extremely excessive values of phylogenetic signal, high mean species' extinction probabilities, and extreme (coalescent) tree shapes. Interestingly, we also observed a decline in the rate of increase in the loss of PD at high phylogenetic clustering (λ → 10) of extinction risks. Our results suggest that the interplay between various aspects of tree shape and a predisposition of higher extinction risks in species-poor clades is required to explain the substantial pruning of the Tree of Life.     

Environmental correlates of the Late Quaternary regional extinctions of large and small Palaearctic mammals

Most studies of mammal extinctions during the Pleistocene–Holocene transition explore the relative effects of climate change vs human impacts on these extinctions, but the relative importance of the different environmental factors involved remains poorly understood. Moreover, these studies are strongly biased towards megafauna, which may have been more influenced by human hunting than species of small body size. We examined the potential environmental causes of Pleistocene–Holocene mammal extinctions by linking regional environmental characteristics with the regional extinction rates of large and small mammals in 14 Palaearctic regions. We found that regional extinction rates were larger for megafauna, but extinction patterns across regions were similar for both size groups, emphasizing the importance of environmental change as an extinction factor as opposed to hunting. Still, the bias towards megafauna extinctions was larger in southern Europe and smaller in central Eurasia. The loss of suitable habitats, low macroclimatic heterogeneity within regions and an increase in precipitation were identified as the strongest predictors of regional extinction rates. Suitable habitats for many species of the Last Glacial fauna were grassland and desert, but not tundra or forest. The low‐extinction regions identified in central Eurasia are characterized by the continuous presence of grasslands and deserts until the present. In contrast, forest expansion associated with an increase in precipitation and temperature was likely the main factor causing habitat loss in the high‐extinction regions. The shift of grassland into tundra also contributed to the loss of suitable habitats in northern Eurasia. Habitat loss was more strongly related to the extinctions of megafauna than of small mammals. Ungulate species with low tolerance to deep snow were more likely to go regionally extinct. Thus, the increase in precipitation at the Pleistocene–Holocene transition may have also directly contributed to the extinctions by creating deep snow cover which decreases forage availability in winter.     

Re-assessing current extinction rates

There is a widespread belief that we are experiencing a mass extinction event similar in severity to previous mass extinction events in the last 600 million years where up to 95% of species disappeared. This paper reviews evidence for current extinctions and different methods of assessing extinction rates including species–area relationships and loss of tropical forests, changing threat status of species, co-extinction rates and modelling the impact of climate change. For 30 years some have suggested that extinctions through tropical forest loss are occurring at a rate of up to 100 species a day and yet less than 1,200 extinctions have been recorded in the last 400 years. Reasons for low number of identified global extinctions are suggested here and include success in protecting many endangered species, poor monitoring of most of the rest of species and their level of threat, extinction debt where forests have been lost but species still survive, that regrowth forests may be important in retaining ‘old growth’ species, fewer co-extinctions of species than expected, and large differences in the vulnerability of different taxa to extinction threats. More recently, others have suggested similar rates of extinction to earlier estimates but with the key cause of extinction being climate change, and in particular rising temperatures, rather than deforestation alone. Here I suggest that climate change, rather than deforestation is likely to bring about such high levels of extinction since the impacts of climate change are local to global and that climate change is acting synergistically with a range of other threats to biodiversity including deforestation.     

Activity restriction and the mechanistic basis for extinctions under climate warming

Correlative analyses predict that anthropogenic climate warming will cause widespread extinction but the nature and generality of the underlying mechanisms is unclear. Warming‐induced activity restriction has been proposed as a general explanatory mechanism for recent population extinctions in lizards, and has been used to forecast future extinction. Here, I test this hypothesis using globally applied biophysical calculations of the effects of warming and shade reduction on potential activity time and whole‐life‐cycle energy budgets. These ‘thermodynamic niche’ analyses show that activity restriction from climate warming is unlikely to provide a general explanation of recent extinctions, and that loss of shade is viable alternative explanation. Climate warming could cause population declines, even under increased activity potential, through joint impacts on fecundity and mortality rates. However, such responses depend strongly on behaviour, habitat (shade, food) and life history, all of which should be explicitly incorporated in mechanistic forecasts of extinction risk under climate change.     

The origin and evolution of coral species richness in a marine biodiversity hotspot*

The Coral Triangle (CT) region of the Indo‐Pacific realm harbors an extraordinary number of species, with richness decreasing away from this biodiversity hotspot. Despite multiple competing hypotheses, the dynamics underlying this regional diversity pattern remain poorly understood. Here, we use a time‐calibrated evolutionary tree of living reef coral species, their current geographic ranges, and model‐based estimates of regional rates of speciation, extinction, and geographic range shifts to show that origination rates within the CT are lower than in surrounding regions, a result inconsistent with the long‐standing center of origin hypothesis. Furthermore, endemism of coral species in the CT is low, and the CT endemics are older than relatives found outside this region. Overall, our model results suggest that the high diversity of reef corals in the CT is largely due to range expansions into this region of species that evolved elsewhere. These findings strongly support the notion that geographic range shifts play a critical role in generating species diversity gradients. They also show that preserving the processes that gave rise to the striking diversity of corals in the CT requires protecting not just reefs within the hotspot, but also those in the surrounding areas.     

Response of Pleistocene Coral Reefs to Environmental Change Over Long Temporal Scales

SYNOPSIS. TWO studies from the Pleistocene coral reef fossilrecord demonstrate the sensitivity of reef communities to bothlocal environmental parameters and habitat reduction. In thefirst study, Pleistocene reef coral assemblages from Papua NewGuinea show pronounced constancy in taxonomic composition andspecies diversity between 125 and 30 ka (thousand years). Spatialdifferences in reef coral community composition during successivehigh stands of sea level were greater among sites of the sameage than among reefs of different ages, even though global changesin sea level, atmospheric CO₂ concentration, tropical benthichabitat area, and temperature varied at each high sea levelstand. Thus, local environmental variation associated with runofffrom the land had greater influence on reef coral communitycomposition than variation in global climate and sea level.Proportional sampling from a regional species pool does notexplain the temporal persistence and local factors likely playeda major role. Examination of coral reef response to global changeshould not only involve regional diversity patterns but alsolocal ecological factors, and the interactive effects of localand global environmental change. In the second study, Pleistocene extinction of two widespread,strictly insular species of Caribbean reef corals, Pocilloporacf. palmata (Geister, 1975) and an organ-pipe growth form ofthe Montastraea "annularis" species complex, was natural anddid not involve gradual decrease in range and abundance, butwas sudden (thousands of years) throughout the entire range.One explanation is that sea level drop at the Last Glacial Maximum(LGM—18 ka) resulted in a threshold of habitat reduction,and caused disruption of coral metapopulation structure. Thresholdeffects predicted by metapopulation dynamics may also explainthe apparent paradox of the large amount of degraded modernreef habitat without any known modern-day reef coral extinctions.The rapid extinction of widespread Pleistocene species emphasizesthe vulnerability of reef corals in the face of present rapidenvironmental and climatic change.     

Climate change effects on animal and plant phylogenetic diversity in southern Africa

Much attention has been paid to the effects of climate change on species' range reductions and extinctions. There is however surprisingly little information on how climate change driven threat may impact the tree of life and result in loss of phylogenetic diversity (PD). Some plant families and mammalian orders reveal nonrandom extinction patterns, but many other plant families do not. Do these discrepancies reflect different speciation histories and does climate induced extinction result in the same discrepancies among different groups? Answers to these questions require representative taxon sampling. Here, we combine phylogenetic analyses, species distribution modeling, and climate change projections on two of the largest plant families in the Cape Floristic Region (Proteaceae and Restionaceae), as well as the second most diverse mammalian order in Southern Africa (Chiroptera), and an herbivorous insect genus (Platypleura) in the family Cicadidae to answer this question. We model current and future species distributions to assess species threat levels over the next 70 years, and then compare projected with random PD survival. Results for these animal and plant clades reveal congruence. PD losses are not significantly higher under predicted extinction than under random extinction simulations. So far the evidence suggests that focusing resources on climate threatened species alone may not result in disproportionate benefits for the preservation of evolutionary history.     

The form of direct interspecific competition modifies secondary extinction patterns in multi‐trophic food webs

Forcibly removing species from ecosystems has important consequences for the remaining assemblage, leading to changes in community structure, ecosystem functioning and secondary (cascading) extinctions. One key question that has arisen from single‐ and multi‐trophic ecosystem models is whether the secondary extinctions that occur within competitive communities (guilds) are also important in multi‐trophic ecosystems? The loss of consumer–resource links obviously causes secondary extinction of specialist consumers (topological extinctions), but the importance of secondary extinctions in multi‐trophic food webs driven by direct competitive exclusion remains unknown. Here I disentangle the effects of extinctions driven by basal competitive exclusion from those caused by trophic interactions in a multi‐trophic ecosystem (basal producers, intermediate and top consumers). I compared food webs where basal species either show diffuse (all species compete with each other identically: no within guild extinctions following primary extinction) or asymmetric competition (unequal interspecific competition: within guild extinctions are possible). Basal competitive exclusion drives extra extinction cascades across all trophic levels, with the effect amplified in larger ecosystems, though varying connectance has little impact on results. Secondary extinction patterns based on the relative abundance of the species lost in the primary extinction differ qualitatively between diffuse and asymmetric competition. Removing asymmetric basal species with low (high) abundance triggers fewer (more) secondary extinctions throughout the whole food web than removing diffuse basal species. Rare asymmetric competitors experience less pressure from consumers compared to rare diffuse competitors. Simulations revealed that diffuse basal species are never involved in extinction cascades, regardless of the trophic level of a primary extinction, while asymmetric competitors were. This work highlights important qualitative differences in extinction patterns that arise when different assumptions are made about the form of direct competition in multi‐trophic food webs.     

How global extinctions impact regional biodiversity in mammals

Phylogenetic diversity (PD) represents the evolutionary history of a species assemblage and is a valuable measure of biodiversity because it captures not only species richness but potentially also genetic and functional diversity. Preserving PD could be critical for maintaining the functional integrity of the world's ecosystems, and species extinction will have a large impact on ecosystems in areas where the ecosystem cost per species extinction is high. Here, we show that impacts from global extinctions are linked to spatial location. Using a phylogeny of all mammals, we compare regional losses of PD against a model of random extinction. At regional scales, losses differ dramatically: several biodiversity hotspots in southern Asia and Amazonia will lose an unexpectedly large proportion of PD. Global analyses may therefore underestimate the impacts of extinction on ecosystem processes and function because they occur at finer spatial scales within the context of natural biogeography.     

Mass extinctions and ocean acidification: biological constraints on geological dilemmas

The five mass extinction events that the earth has so far experienced have impacted coral reefs as much or more than any other major ecosystem. Each has left the Earth without living reefs for at least four million years, intervals so great that they are commonly referred to as ‘reef gaps’ (geological intervals where there are no remnants of what might have been living reefs). The causes attributed to each mass extinction are reviewed and summarised. When these causes and the reef gaps that follow them are examined in the light of the biology of extant corals and their Pleistocene history, most can be discarded. Causes are divided into (1) those which are independent of the carbon cycle: direct physical destruction from bolides, ‘nuclear winters’ induced by dust clouds, sea-level changes, loss of area during sea-level regressions, loss of biodiversity, low and high temperatures, salinity, diseases and toxins and extraterrestrial events and (2) those linked to the carbon cycle: acid rain, hydrogen sulphide, oxygen and anoxia, methane, carbon dioxide, changes in ocean chemistry and pH. By process of elimination, primary causes of mass extinctions are linked in various ways to the carbon cycle in general and ocean chemistry in particular with clear association with atmospheric carbon dioxide levels. The prospect of ocean acidification is potentially the most serious of all predicted outcomes of anthropogenic carbon dioxide increase. This study concludes that acidification has the potential to trigger a sixth mass extinction event and to do so independently of anthropogenic extinctions that are currently taking place.     

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.