Permian–Triassic Osteichthyes (bony fishes): diversity dynamics and body size evolution

The Permian and Triassic were key time intervals in the history of life on Earth. Both periods are marked by a series of biotic crises including the most catastrophic of such events, the end‐Permian mass extinction, which eventually led to a major turnover from typical Palaeozoic faunas and floras to those that are emblematic for the Mesozoic and Cenozoic. Here we review patterns in Permian–Triassic bony fishes, a group whose evolutionary dynamics are understudied. Based on data from primary literature, we analyse changes in their taxonomic diversity and body size (as a proxy for trophic position) and explore their response to Permian–Triassic events. Diversity and body size are investigated separately for different groups of Osteichthyes (Dipnoi, Actinistia, ‘Palaeopterygii’, ‘Subholostei’, Holostei, Teleosteomorpha), within the marine and freshwater realms and on a global scale (total diversity) as well as across palaeolatitudinal belts. Diversity is also measured for different palaeogeographical provinces. Our results suggest a general trend from low osteichthyan diversity in the Permian to higher levels in the Triassic. Diversity dynamics in the Permian are marked by a decline in freshwater taxa during the Cisuralian. An extinction event during the end‐Guadalupian crisis is not evident from our data, but ‘palaeopterygians’ experienced a significant body size increase across the Guadalupian–Lopingian boundary and these fishes upheld their position as large, top predators from the Late Permian to the Late Triassic. Elevated turnover rates are documented at the Permian–Triassic boundary, and two distinct diversification events are noted in the wake of this biotic crisis, a first one during the Early Triassic (dipnoans, actinistians, ‘palaeopterygians’, ‘subholosteans’) and a second one during the Middle Triassic (‘subholosteans’, neopterygians). The origination of new, small taxa predominantly among these groups during the Middle Triassic event caused a significant reduction in osteichthyan body size. Neopterygii, the clade that encompasses the vast majority of extant fishes, underwent another diversification phase in the Late Triassic. The Triassic radiation of Osteichthyes, predominantly of Actinopterygii, which only occurred after severe extinctions among Chondrichthyes during the Middle–Late Permian, resulted in a profound change within global fish communities, from chondrichthyan‐rich faunas of the Permo‐Carboniferous to typical Mesozoic and Cenozoic associations dominated by actinopterygians. This turnover was not sudden but followed a stepwise pattern, with leaps during extinction events.     

Climatic drivers of latitudinal variation in Late Triassic tetrapod diversity

The latitudinal biodiversity gradient (LBG), the increase in biodiversity from the poles to the equator, is one of the most widely recognized global macroecological patterns, yet its deep time evolution and drivers remain uncertain. The Late Triassic (237–201 Ma), a critical interval for the early evolution and radiation of modern tetrapod groups (e.g. crocodylomorphs, dinosaurs, mammaliamorphs), offers a unique opportunity to explore the palaeolatitudinal patterns of tetrapod diversity since it is extensively sampled spatially when compared with other pre‐Cenozoic intervals, particularly at lower palaeolatitudes. Here, we explore palaeolatitudinal patterns of Late Triassic tetrapod diversity by applying sampling standardization to comprehensive occurrence data from the Paleobiology Database (PBDB). We then use palaeoclimatic model simulations to explore the palaeoclimatic ranges occupied by major tetrapod groups, allowing insight into the influence of palaeoclimate on the palaeolatitudinal distribution of these groups. Our results show that Late Triassic tetrapods generally do not conform to a modern‐type LBG; instead, sampling‐standardized species richness is highest at mid‐palaeolatitudes. In contrast, the richness of pseudosuchians (crocodylians and their relatives) is highest at the palaeoequator, a pattern that is retained throughout their subsequent evolutionary history. Pseudosuchians generally occupied a more restricted range of palaeoclimatic conditions than other tetrapod groups, a condition analogous to modern day reptilian ectotherms, while avemetatarsalians (the archosaur group containing dinosaurs and pterosaurs) exhibit comparatively wider ranges, which is more similar to modern endotherms, such as birds and mammals, suggesting important implications for the evolution of thermal physiology in dinosaurs.     

Global patterns and predictors of tropical reef fish species richness

In the marine realm, the tropics host an extraordinary diversity of taxa but the drivers underlying the global distribution of marine organisms are still under scrutiny and we still lack an accurate global predictive model. Using a spatial database for 6336 tropical reef fishes, we attempted to predict species richness according to geometric, biogeographical and environmental explanatory variables. In particular, we aimed to evaluate and disentangle the predictive performances of temperature, habitat area, connectivity, mid‐domain effect and biogeographical region on reef fish species richness. We used boosted regression trees, a flexible machine‐learning technique, to build our predictive model and structural equation modeling to test for potential ‘mediation effects’ among predictors. Our model proved to be accurate, explaining 80% of the total deviance in fish richness using a cross‐validated procedure. Coral reef area and biogeographical region were the primary predictors of reef fish species richness, followed by coast length, connectivity, mid‐domain effect and sea surface temperature, with interactions between the region and other predictors. Important indirect effects of water temperature on reef fish richness, mediated by coral reef area, were also identified. The relationship between environmental predictors and species richness varied markedly among biogeographical regions. Our analysis revealed that a few easily accessible variables can accurately predict reef fish species richness. They also highlight concerns regarding ongoing environmental declines, with region‐specific responses to variation in environmental conditions predicting a variable response to anthropogenic impacts.     

Oxygen‐restricted facies of the basal Jurassic of North West Europe

Events across the Triassic‐Jurassic boundary are a matter of great interest because this marks the time of one of the five biggest marine mass extinctions in the Phanerozoic. Some of the best sections across the boundary are in north west Europe; attention here is focussed on the basal Jurassic. Sedimentological and palaeoecological study has been undertaken of a number of sections in England and Germany, most of them borehole cores, and indicate a range of oxygen‐restricted facies, signifying episodic fluctuations between anoxic and dysoxic conditions. Analysis of the Th/U ratio in the coastal outcrop of St. Audrie's, Somerset, confirms this interpretation. The Triassic‐Jurassic boundary in north west Europe is characterised by a regression‐transgression couplet, with the corresponding sea‐level change being quite possibly of short duration.     

‘Fish’ (Actinopterygii and Elasmobranchii) diversification patterns through deep time

Actinopterygii (ray‐finned fishes) and Elasmobranchii (sharks, skates and rays) represent more than half of today's vertebrate taxic diversity (approximately 33000 species) and form the largest component of vertebrate diversity in extant aquatic ecosystems. Yet, patterns of ‘fish’ evolutionary history remain insufficiently understood and previous studies generally treated each group independently mainly because of their contrasting fossil record composition and corresponding sampling strategies. Because direct reading of palaeodiversity curves is affected by several biases affecting the fossil record, analytical approaches are needed to correct for these biases. In this review, we propose a comprehensive analysis based on comparison of large data sets related to competing phylogenies (including all Recent and fossil taxa) and the fossil record for both groups during the Mesozoic–Cainozoic interval. This approach provides information on the ‘fish’ fossil record quality and on the corrected ‘fish’ deep‐time phylogenetic palaeodiversity signals, with special emphasis on diversification events. Because taxonomic information is preserved after analytical treatment, identified palaeodiversity events are considered both quantitatively and qualitatively and put within corresponding palaeoenvironmental and biological settings. Results indicate a better fossil record quality for elasmobranchs due to their microfossil‐like fossil distribution and their very low diversity in freshwater systems, whereas freshwater actinopterygians are diverse in this realm with lower preservation potential. Several important diversification events are identified at familial and generic levels for elasmobranchs, and marine and freshwater actinopterygians, namely in the Early–Middle Jurassic (elasmobranchs), Late Jurassic (actinopterygians), Early Cretaceous (elasmobranchs, freshwater actinopterygians), Cenomanian (all groups) and the Paleocene–Eocene interval (all groups), the latter two representing the two most exceptional radiations among vertebrates. For each of these events along with the Cretaceous‐Paleogene extinction, we provide an in‐depth review of the taxa involved and factors that may have influenced the diversity patterns observed. Among these, palaeotemperatures, sea‐levels, ocean circulation and productivity as well as continent fragmentation and environment heterogeneity (reef environments) are parameters that largely impacted on ‘fish’ evolutionary history, along with other biotic constraints.     

Facies and geochemical evidence bearing on the end‐triassic disappearance of the alpine reef ecosystem

A carbon and oxygen isotope profile is presented across the Triassic‐Jurassic boundary at the classic locality of Kendelbach, Austria. In conjunction with faciès data it lends no support to the claim that the spectacular disappearance of the reef ecosystem at the end of the Triassic was due to a sharp fall of seawater temperature. A model relating mass extinction to sea‐level change is preferred.     

Diversity patterns in upper Cambrian to Lower Ordovician trilobite communities of north‐western Argentina

The hierarchical structure of biodiversity from a regional scale analysis has received much attention as an alternative approach to unravelling the principal drivers of biodiversification. To better understand the processes that control the diversification of Cambro‐Ordovician trilobite communities from the Argentine Cordillera Oriental, we explore patterns of occupancy and diversity trajectories at the local and regional scales through seven intervals (Furongian, loTr1, upTr1, loTr2, upTr2, Tr3 and Fl2–3), and across an onshore‐offshore profile. Our results indicate: (1) a decrease in regional diversity from the upper Tr2 onwards, mainly caused by a reduction in the number of rare taxa, coupled with stable beta diversity at regional scale and a constant rise in beta diversity in deep subtidal environments; (2) a higher proportion of regional diversity allocated to the within‐habitat beta component; and (3) that changes in gamma diversity are driven primarily by changes in alpha diversity during the Furongian–Tr3, whereas in the Floian, beta diversity seems to modulate regional diversity. These trends and associated patterns indicate increasing ecological differences among taxa, shifting from metacommunities where most taxa have similar ecological preferences or ‘Hubbell type’ to metacommunities with high niche differentiation or ‘Hutchinson type’. Interestingly, the timing of this shift coincides with the regional‐scale turnover between trilobite evolutionary faunas suggesting that the rise in niche differentiation among these genera may be related to the transition. Superimposed on this general trend, particular diversity structures can be understood in the light of metacommunity dynamics, such as dispersal limitation and mass effect.     

The effect of habitat on modern shark diversification

Sharks occupy marine habitats ranging from shallow, inshore environments to pelagic, and deepwaters, and thus provide a model system for testing how gross habitat differences have shaped vertebrate macroevolution. Palaeontological studies have shown that onshore lineages diversify more quickly than offshore taxa. Among onshore habitats, coral reef‐association has been shown to increase speciation rates in several groups of fishes and invertebrates. In this study, we investigated whether speciation rates are habitat dependent by generating the first comprehensive molecular timescale for shark divergence. Using phylogenetic comparative methods, we rejected the hypothesis that shelf (i.e. onshore) lineages have higher speciation rates compared to those occupying deepwater and oceanic (i.e. offshore) habitats. Our results, however, support the hypothesis of increased speciation rates in coral reef‐associated lineages within the Carcharhinidae. Our new timetree suggests that the two major shark lineages leading to the extant shark diversity began diversifying mostly after the end‐Permian mass extinction: the squalimorphs into deepwater and the galeomorphs into shelf habitats. We suggest that the breakdown of the onshore–offshore speciation rate pattern in sharks is mediated by success in deepwater environments through ecological partitioning, and in some cases, the evolution of morphological novelty.     

The latitudinal position of peak marine diversity in living and fossil biotas

Aim Peak marine taxonomic diversity has only rarely occurred at or near the equator during the Phanerozoic Eon, in contrast to the present‐day pattern. This fundamental difference is difficult to reconcile because the latitude at which peak diversity occurs for living marine taxa has not yet been explicitly determined at a broad taxonomic and spatial scale. Here, we attempt to determine this value in order to compare the contemporary and fossil patterns directly. Location Our data are global in coverage. Methods We used a literature compilation of 149 present‐day marine latitudinal diversity gradients. We summed the number of marine taxa that exhibited peak diversity within 10° latitudinal bins. In addition, we recorded locality data, general habitat (benthic/pelagic), and the taxonomic level of the study organisms. Results We found that peak diversity for most sampled marine taxa currently occurs between 10° and 20° N, even after correcting for a Northern Hemisphere sampling bias. Moreover, this peak position is a global phenomenon: it is found across habitats and higher taxa, within all sampled ocean basins, and on both sides of the Atlantic and Pacific oceans. Benthic taxa, which dominate our data, exhibit one peak at 10°–20° N, while pelagic taxa exhibit a peak at 10°–20° N and an additional peak at 10°–20° S, producing a distinct trough at the equator. Main conclusions Our data indicate that peak marine diversity for many taxa is currently within 10°–20° N rather than at the equator, and that this is not likely to result from either undersampling at lower latitudes or the pattern being dominated by a particular taxon. Possible explanations may include a coincidence with the intertropical convergence zone, a mid‐domain effect, abundant shallow marine habitat, or high ocean temperatures at latitudes nearest the equator. Regardless of its exact cause, the position of peak diversity should be considered a fundamental feature of the latitudinal diversity gradient that must be accounted for within attempts to explain the latter’s existence.     

Dynamics of taxonomic diversity of bivalves in the Phanerozoic

Changes in the taxonomic composition of bivalves during the Phanerozoic are considered. Each period is characterized by a special set of taxa, in particular, families. Changes in taxonomic diversity, the episodes of maximum and minimum diversity are established and compared with those of other invertebrate groups. In general, the taxonomic diversity of bivalves gradually increased, except for a sharp decrease in the Early Triassic.     

Recovery of benthic marine communities from the end‐Permian mass extinction at the low latitudes of eastern Panthalassa

Based on the quantitative community analysis using species‐level identifications, we track the restoration of benthic ecosystems after the end‐Permian mass extinction throughout the Lower Triassic of the western USA. New data on the palaeoecology of the Thaynes Group and Sinbad Formation are provided, which fill a gap between the recently studied palaeoecology of the Griesbachian–Dienerian Dinwoody Formation and the Spathian Virgin Formation. In the Sinbad Formation and Thaynes Group, 17 species (12 genera) of bivalves, 7 species and genera of gastropods and 2 species and genera of brachiopods are recognized. The new bivalve genus Confusionella (Pteriidae) is described. A comprehensive review of the whole Lower Triassic succession of benthic ecosystems of the western USA indicates that mid‐ and inner shelf environments show incipient recovery signals around the Griesbachian–Dienerian transition, during the Smithian and, most profound, during the early Spathian. Ecological data from youngest strata of the Dinwoody Formation as well as stratigraphic ranges of species suggest that the late Dienerian was likely a time interval of environmental stress for benthic ecosystems. Despite some evidence for short‐term environmental disturbances (e.g. shift of dominant taxa, transient drop in alpha‐diversity) during the Smithian–Spathian transition, benthic ecosystems did not show any notable taxonomic turnover at that time, in contrast to the major crisis that affected ammonoids and conodonts. Whereas alpha‐diversity of benthic communities generally increased throughout the Early Triassic, beta‐diversity remained low, which reflects a persistently wide environmental range of benthic species. This observation is in accordance with a recently proposed model that predicts a time lag between increasing within‐habitat diversity (alpha‐diversity) and the onset of taxonomic differentiation between habitats (beta‐diversity) during biotic recoveries from mass extinction events. The observation that beta‐diversity had not significantly increased during the Early Triassic might also provide an explanation for the comparably sluggish increase in benthic diversity during that time, which has previously been attributed to persistent environmental stress.     

Harnessing stratigraphic bias at the section scale: conodont diversity in the Homerian (Silurian) of the Midland Platform,England

Fossil abundance and diversity in geological successions are subject to bias arising from shifting depositional and diagenetic environments, resulting in variable rates of fossil accumulation and preservation. In simulations, this bias can be constrained based on sequence‐stratigraphic architecture. Nonetheless, a practical quantitative method of incorporating the contribution of sequence‐stratigraphic architecture in community palaeoecology and diversity analyses derived from individual successions is missing. As a model of faunal turnover affected by the stratigraphic bias, we use the ‘Mulde event’, a postulated mid‐Silurian interval of elevated conodont turnover, which coincides with global eustatic sea‐level changes and which has been based on regionally constrained observations. We test whether conodont turnover is highest at the boundary corresponding to the ‘event’ and post‐‘event’ interval against the alternative that conodont turnover reflects habitat tracking and peaks at facies shifts. Based on the previously documented, parasequence‐level stratigraphic framework of sections in the northern and central part of the Midland Platform, the relative controls of sequence‐stratigraphic architecture, time and depositional environment over conodont distribution are evaluated using permutational multivariate analysis of variance. The depositional environment controls the largest part of variability in conodont assemblage composition, whereas the postulated ‘Mulde event’, or genuine temporal change in conodont diversity, cannot be detected. Depending on the binning of the stratigraphic succession, contrasting diversity and turnover patterns can be produced. The simple approach proposed here, emulating partitioning of β diversity into spatial and temporal components, may help to constrain the stratigraphic bias, even at the scale of an individual section.     

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.     

Fossil insects of the middle and upper Permian of European Russia

Fossil insects of European Russia from the Urzhumian to Vyatkian stages are reviewed, new taxa are described, and dynamics of insect taxonomic diversity around the Permian-Triassic boundary in light of the Paleozoic-Mesozoic boundary global extinction problem is analyzed. Traces of interactions between arthropods and plants are analyzed. Insect-bearing deposits of the Late Paleozoic found in the northern and eastern areas of the East European Platform are unique on the global scale in their completeness and continuity, allowing us to trace especially comprehensively the biotic processes that occurred around the boundary described as the time of the greatest biotic catastrophe of the Phanerozoic. A total of 28 genera and 111 species are newly described. Within the range from the Urzhumian to the Permo-Triassic boundary, 15 representative successive assemblages, including 112 families, are recognized (seven in the area in question and eight in other regions of Asia, Australia, and Africa). New tools are developed for the analysis of the dynamics of diversity. These tools show an approximately equilibrium (slightly positive) dynamics in the Urzhumian and Severodvinian and a drop in diversity during the Vyatkian Age. It is shown that Permian insect assemblages acquired a substantially post-Paleozoic pattern much earlier than the end of the Paleozoic. The character of changes that took place in the Induan and Olenekian remains uncertain, but a large-scale extinction event did not occur here: most families that have not been recorded at the beginning of the Triassic are recorded again in the Middle and Upper Triassic. Nevertheless, a biotic crisis probably actually took place, but was reduced to reorganization of the biota’s structure, which provided enormous growth of biodiversity over subsequent hundreds of millions of years, rather than resulted in catastrophic extinction. This study is intended for entomologists, stratigraphers, and all readers interested in the biotic events that took place around the Permian-Triassic boundary.     

Galvechelone lopezmartinezae gen. et sp. nov., a new cryptodiran turtle in the Lower Cretaceous of Europe

Abstract: The Spanish town of Galve (Teruel) is notable because of the abundance of Upper Jurassic and, especially, Lower Cretaceous vertebrates recorded there. Although most groups have been studied in detail, information on turtles is very limited even though the material is relatively abundant. So far, no turtle taxa have been identified at the generic level. The only Lower Cretaceous articulated specimen from Galve is analysed here. It is identified as a representative of Cryptodira, Galvechelone lopezmartinezae gen. et sp. nov. Galvechelone lopezmartinezae is determined as a taxon belonging to the node that groups the turtles traditionally assigned to ‘Macrobaenidae’ and ‘Sinemydidae’, and other taxa such as the members of Panchelonioidea. This node, very abundant in the Lower Cretaceous of Asia, and with a broad subsequent distribution, has recently been recognized in the Lower Cretaceous of Europe. The diversity of basal members of Eucryptodira in the European Late Jurassic (represented by Thalassemydidae, Plesiochelyidae and Eurysternidae) was high. Owing to a relative scarcity of well‐preserved early Cretaceous turtles from Europe, the knowledge of this group of reptiles is limited. The study of the new turtle from Galve, together with the recently described Hoyasemys jimenezi, and the recently completed review of the enigmatic Chitracephalus dumonii demonstrate that members of the cryptodiran node grouping ‘Macrobaenidae’, ‘Sinemydidae’ and Panchelonioidea were also very diverse in this period.     

Escaping the heat: range shifts of reef coral taxa in coastal Western Australia

One of the most critical challenges facing ecologists today is to understand the changing geographic distribution of species in response to current and predicted global warming. Coastal Western Australia is a natural laboratory in which to assess the effect of climate change on reef coral communities over a temporal scale unavailable to studies conducted solely on modern communities. Reef corals composing Late Pleistocene reef assemblages exposed at five distinct localities along the west Australian coast were censused and the results compared with coral occurrence data published for the modern reefs offshore of each locality. The resulting comparative data set comprises modern and Late Pleistocene reef coral communities occurring over approximately 12° of latitude. For the modern reefs this gradient includes the zone of overlap between the Dampierian and Flindersian Provinces. Modern reef coral communities show a pronounced gradient in coral composition over the latitudinal range encompassed by the study, while the gradient in community composition is not as strong for Pleistocene communities. Tropical‐adapted taxa contracted their ranges north since Late Pleistocene time, emplacing two biogeographic provinces in a region in which a single province had existed previously. Beta diversity values for adjacent communities also reflect this change. Modern reefs show a distinct peak in beta diversity in the middle of the region; the peak is not matched by Pleistocene reefs. Beta diversity is correlated with distance only for comparisons between modern reefs in the north and the fossil assemblages, further supporting change in distribution of the biogeographic provinces in the study area. Coral taxa present in modern communities clearly expanded and contracted their geographic ranges in response to climate change. Those taxa that distinguish Pleistocene from modern reefs are predicted to migrate south in response to future climate change, and potentially persist in ‘temperature refugia’ as tropical reef communities farther north decline.     

Evolutionary dispersal drives the latitudinal diversity gradient of stony corals

The diversity of stony corals displays one of the most exemplary latitudinal gradients on the planet, yet the evolutionary dynamics that produced this pattern remains unclear. Using both paleontological and distributional data, we compare the origination, extinction and immigration levels between low and high latitudes since the earliest proliferation of the group during the mid‐Triassic. Altogether, first and last occurrence localities in the fossil record do not support a positive preference towards either latitudinal bin. Nonetheless, considering past and present scleractinian fauna, the process of extinction has been apparently more pronounced at higher latitudes based on face values and correlation coefficients. Far above these differences, immigration of extant taxa has been substantially higher towards the tropics than to temperate regions. While the net dispersal toward low latitudes persists in all temporal intervals, the gradient of diversity was largely built up during the Cenozoic Era and only becomes significantly steep from the Neogene Period onwards. This dynamic supports the ‘into the tropical museum’ model, which suggests that tropics have historically acted as a center of accumulation for marine biodiversity.     

The Late Palaeozoic phytoplankton blackout — Artefact or evidence of global change?

The fossil record of the Palaeozoic documents one of the most dramatic changes in Phanerozoic marine primary production, although causes and effects of these changes have been the subject of rather controversial discussions. During the early Palaeozoic the marine phytoplankton experienced an enormous radiation and diversification of taxa especially among acritarchs, which was punctuated by a few extinction events possibly associated with climate changes. Phytoplankton diversity was drastically reduced at the Devonian/Carboniferous boundary, a phenomenon here designated as the “Phytoplankton Blackout”.After a lapse of about 130 million years phytoplankton diversity was gradually restored during the Late Triassic with the appearance of dinoflagellates and somewhat later of coccolithophorids and diatoms. Evidence from recent phytoplankton suggests that the dominant groups of phytoplankton differ from those of the Palaeozoic in their nutritional requirements and that fundamental changes in ocean chemistry have played an important role.The remarkable temporal coincidence of the Phytoplankton Blackout with plate tectonic processes during the assembly and breakup of Pangaea and the concomitant rise of land plants towards the end of the Devonian provide arguments that nutrient depletion in the ocean may have played a decisive role in controlling the phytoplankton blackout. Contrary arguments in favour of oceanic eutrophication, changes in life cycles of dominant phytoplankton groups or selective preservation are discussed and weighed against the scenario presented here. Metazoan evolution seems only loosely linked with the phytoplankton blackout. However, there is good agreement between phytoplankton and reef evolution throughout the Phanerozoic.     

CALIBRATED DIVERSITY,TREE TOPOLOGY AND THE MOTHER OF MASS EXTINCTIONS: THE LESSON OF TEMNOSPONDYLS

Abstract: Three family‐level cladistic analyses of temnospondyl amphibians are used to evaluate the impact of taxonomic rank, tree topology, and sample size on diversity profiles, origination and extinction rates, and faunal turnover. Temnospondyls are used as a case study for investigating replacement of families across the Permo‐Triassic boundary and modality of recovery in the aftermath of the end‐Permian mass extinction. Both observed and inferred (i.e. tree topology‐dependent) values of family diversity have a negligible effect on the shape of the diversity curve. However, inferred values produce both a flattening of the curve throughout the Cisuralian and a less pronounced increase in family diversity from Tatarian through to Induan than do observed values. Diversity curves based upon counts of genera and species display a clearer distinction between peaks and troughs. We use rarefaction techniques (specifically, rarefaction of the number of genera and species within families) to evaluate the effect of sampling size on the curve of estimated family‐level diversity during five time bins (Carboniferous; Cisuralian; Guadalupian–Lopingian; Early Triassic; Middle Triassic–Cretaceous). After applying rarefaction, we note that Cisuralian and Early Triassic diversity values are closer to one another than they are when the observed number of families is used; both values are also slightly higher than the Carboniferous estimated diversity. The Guadalupian–Lopingian value is lower than raw data indicate, reflecting in part the depauperate land vertebrate diversity from the late Cisuralian to the middle Guadalupian (Olson’s gap). The time‐calibrated origination and extinction rate trajectories plot out close to one another and show a peak in the Induan, regardless of the tree used to construct them. Origination and extinction trajectories are disjunct in at least some Palaeozoic intervals, and background extinctions exert a significant role in shaping temnospondyl diversity in the lowermost Triassic. Finally, species‐, genus‐, and family trajectories consistently reveal a rapid increase in temnospondyl diversity from latest Permian to earliest Triassic as well as a decline near the end of the Cisuralian. However, during the rest of the Cisuralian family diversity increases slightly and there is no evidence for a steady decline, contrary to previous reports.     

Additive diversity partitioning in palaeobiology: revisiting Sepkoski’s question

Abstract: Using Whittaker’s concepts of alpha, beta, and gamma diversity, Sepkoski asked how global diversity was assembled at scales ranging from the community to the province. In the years since, ecologists have recast diversity in terms of additive partitions where total diversity can be decomposed into sample‐level alpha diversity plus the sum of a series of beta diversity terms that reflect progressively larger spatial scales. Given that marine alpha diversity represents a tiny fraction of global diversity, Phanerozoic global diversity patterns must be dominated by changes in beta diversity at one or more scales. A ballooning ecological literature demonstrates wide variation in beta diversity among ecosystems, regions, and taxa, suggesting that large changes in beta diversity on evolutionary timescales are likely. But the question is which scales are the most important. Several recent palaeontological studies help to constrain beta diversity within sedimentary basins, and the emergence of sample‐based databases puts an answer to Sepkoski’s question within reach. A new method for calculating diversity partitions for richness is introduced, which allows the calculation of each species’ contribution to alpha and beta diversity, as well as the contribution of each sampling unit to beta diversity.