Global diversity curves reflect more than just the number of taxa that have existed through time: they also mirror variation in the nature of the fossil record and the way the record is reported. These sampling effects are best quantified by assembling and analyzing large numbers of locality-specific biotic inventories. Here, we introduce a new database of this kind for the Phanerozoic fossil record of marine invertebrates. We apply four substantially distinct analytical methods that estimate taxonomic diversity by quantifying and correcting for variation through time in the number and nature of inventories. Variation introduced by the use of two dramatically different counting protocols also is explored. We present sampling-standardized diversity estimates for two long intervals that sum to 300 Myr (Middle Ordovician-Carboniferous; Late Jurassic-Paleogene). Our new curves differ considerably from traditional, synoptic curves. For example, some of them imply unexpectedly low late Cretaceous and early Tertiary diversity levels. However, such factors as the current emphasis in the database on North America and Europe still obscure our view of the global history of marine biodiversity. These limitations will be addressed as the database and methods are refined.

The latitudinally diachronous appearance of angiosperm pollen during the Cretaceous is well documented, but the subsequent diversification and accompanying significant changes in floristic dominance have not been assessed quantitatively for a wide range of paleolatitudes. Trend surfaces fitted to within-palynoflora diversity data from 1125 pollen and spore assemblages show that angiosperms first become floristically prominent in low paleolatitude areas( approximately 20 degrees N to 20 degrees S). Non-magnoliid dicotyledons show a similar but slightly delayed pattern of increase and are the principal component of angiosperm diversity from all areas sampled. Monocotyledons and magnoliid dicotyledons are significant primarily in low to middle paleolatitude palynofloras( approximately 50 degrees N to 20 degrees S) during the latest Cretaceous. As angiosperms become increasingly prevalent the importance of most non-angiosperm taxa either decreases or remains unchanged. The only apparent exception is a striking increase in gnetalean diversity concurrent with the initial angiosperm diversification at low paleolatitudes.

The Cretaceous radiation of angiosperms initiated a major reorganization of terrestrial plant communities as dominance by pteridophytic and gymnospermic groups eventually gave way to dominance by angiosperms. Previously, patterns of biotic replacement have been assessed using measures based on taxonomic diversity data. However, using measures of both abundance and diversity to investigate replacement patterns provides more information about macroecological change in the fossil record than either can provide alone. Analyses of an updated and expanded database of North American palynological samples from Cretaceous sediments document a rapid increase in angiosperm diversity and abundance within individual fossil palynofloras (represent ing local/subregional vegetation). New analyses of floristic diversity patterns support previous results and indicate that the decline of free-sporing plants is more pronounced than the decline of gymnosperms. In contrast, analyses of abundance data appear to show that the decline of gymnosperms is far more pronounced than the decline of free-sporing plants. Detailed examination of both data sets segregated by paleolatitude shows that this apparent contradiction reflects biogeographical differences in the patterns of vegetational change (e.g., free-sporing plants declined in abundance at lower latitudes) as well as sampling bias (e.g., greater sampling in the northern region in the Late Cretaceous). Analyses accounting for these biases support the conclusion that as an giosperms radiated, free-sporing plants rather than gymnosperms (in this case, mainly conifers) experienced the most pronounced decline. A thorough understanding of the Cretaceous radiation of angiosperms will require both abundance and diversity data. It also will require expanding the analyses presented here into other geographic regions as well as sampling more completely at all spatial scales.

Previous analyses of evolutionary patterns, or modes, in fossil lineages have focused overwhelmingly on three simple models: stasis, random walks, and directional evolution. Here we use likelihood methods to fit an expanded set of evolutionary models to a large compilation of ancestor-descendant series of populations from the fossil record. In addition to the standard three models, we assess more complex models with punctuations and shifts from one evolutionary mode to another. As in previous studies, we find that stasis is common in the fossil record, as is a strict version of stasis that entails no real evolutionary changes. Incidence of directional evolution is relatively low (13%), but higher than in previous studies because our analytical approach can more sensitively detect noisy trends. Complex evolutionary models are often favored, overwhelmingly so for sequences comprising many samples. This finding is consistent with evolutionary dynamics that are, in reality, more complex than any of the models we consider. Furthermore, the timing of shifts in evolutionary dynamics varies among traits measured from the same series. Finally, we use our empirical collection of evolutionary sequences and a long and highly resolved proxy for global climate to inform simulations in which traits adaptively track temperature changes over time. When realistically calibrated, we find that this simple model can reproduce important aspects of our paleontological results. We conclude that observed paleontological patterns, including the prevalence of stasis, need not be inconsistent with adaptive evolution, even in the face of unstable physical environments.evolutionary mode | stasis | gradualism | punctuated equilibrium | climate change P aleontologists have long sought to document patterns of trait change within fossil species and to infer from these patterns their underlying evolutionary drivers (1-3). However, only recently have sufficient case studies accumulated to assess what the aggregated fossil record has to say about phenotypic evolution occurring on the 10 5 -to 10 7 -y timescales routinely captured in paleontological sequences. Several factors have contributed to this cumulative increase, including advances in geochronology that more readily permit time-calibrating evolutionary sequences, better morphometric practices for documenting trait change, and new analytical tools to examine the resulting data. Perhaps most important, however, is that paleontologists were motivated to capture many additional examples of trait evolution in fossil lineages in response to the intense debate centered around punctuated equilibrium (4-6)-the notion that species do not usually evolve gradually over long periods of time but instead emerge in a discontinuous pattern, in bursts of change associated with cladogenesis followed by longer intervals of morphological stasis.Punctuated equilibrium proponents and critics disagreed about how best to interpret the same patterns of trait evolution, and thus early reviews (7, 8) did little to reso...

Matrotrophy, the continuous extra‐vitelline supply of nutrients from the parent to the progeny during gestation, is one of the masterpieces of nature, contributing to offspring fitness and often correlated with evolutionary diversification. The most elaborate form of matrotrophy—placentotrophy—is well known for its broad occurrence among vertebrates, but the comparative distribution and structural diversity of matrotrophic expression among invertebrates is wanting. In the first comprehensive analysis of matrotrophy across the animal kingdom, we report that regardless of the degree of expression, it is established or inferred in at least 21 of 34 animal phyla, significantly exceeding previous accounts and changing the old paradigm that these phenomena are infrequent among invertebrates. In 10 phyla, matrotrophy is represented by only one or a few species, whereas in 11 it is either not uncommon or widespread and even pervasive. Among invertebrate phyla, Platyhelminthes, Arthropoda and Bryozoa dominate, with 162, 83 and 53 partly or wholly matrotrophic families, respectively. In comparison, Chordata has more than 220 families that include or consist entirely of matrotrophic species. We analysed the distribution of reproductive patterns among and within invertebrate phyla using recently published molecular phylogenies: matrotrophy has seemingly evolved at least 140 times in all major superclades: Parazoa and Eumetazoa, Radiata and Bilateria, Protostomia and Deuterostomia, Lophotrochozoa and Ecdysozoa. In Cycliophora and some Digenea, it may have evolved twice in the same life cycle. The provisioning of developing young is associated with almost all known types of incubation chambers, with matrotrophic viviparity more widespread (20 phyla) than brooding (10 phyla). In nine phyla, both matrotrophic incubation types are present. Matrotrophy is expressed in five nutritive modes, of which histotrophy and placentotrophy are most prevalent. Oophagy, embryophagy and histophagy are rarer, plausibly evolving through heterochronous development of the embryonic mouthparts and digestive system. During gestation, matrotrophic modes can shift, intergrade, and be performed simultaneously. Invertebrate matrotrophic adaptations are less complex structurally than in chordates, but they are more diverse, being formed either by a parent, embryo, or both. In a broad and still preliminary sense, there are indications of trends or grades of evolutionarily increasing complexity of nutritive structures: formation of (i) local zones of enhanced nutritional transport (placental analogues), including specialized parent–offspring cell complexes and various appendages increasing the entire secreting and absorbing surfaces as well as the contact surface between embryo and parent, (ii) compartmentalization of the common incubatory space into more compact and ‘isolated’ chambers with presumably more effective nutritional relationships, and (iii) internal secretory (‘milk’) glands. Some placental analogues in onychophorans and arthropods mimi...

Summary 1. We have examined large‐scale geographical patterns in species richness for continental shelf bryozoan assemblages in the North Atlantic. Bryozoans are common and often abundant benthic organisms, but they have not previously been examined at this scale of resolution. 2. Assemblage species richness was estimated by sample species richness. This was highest at intermediate depths (10–75 m) at all latitudes where there were sufficient data, but there was no statistically significant variation with depth for the overall data set (all latitudes pooled). Mean assemblage species richness showed no significant variation with latitude, although the highest individual values were generally from lower latitudes. There is thus as yet no convincing evidence for a latitudinal cline in the alpha diversity of North Atlantic bryozoans. 3. Pooling of data into bins of 10 degrees of latitude, or into biogeographic provinces, to estimate regional species richness revealed significant undersampling. Two independent techniques to correct for this undersampling revealed a latitudinal cline in the regional species richness of North Atlantic bryozoans, with a peak around 10–30°N, and a steady decrease in richness north to 80°N. 4. Two measures of beta diversity (Whittaker and Jaccard) revealed relatively high turnover, presumably related to habitat heterogeneity within regional bins or to significant environmental variation across bins. There was a tendency for beta diversity to be higher at lower latitudes, as would be expected from a combination of a latitudinal cline in regional diversity with a mean assemblage species richness invariant with latitude. Null models were used to clarify the expected relationship between the two measures of beta diversity, and these indicated a strong influence of species‐abundance structure in the North Atlantic bryozoan data.

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The chemical composition of fossil soft tissues is a potentially powerful and yet underutilized tool for elucidating the affinity of problematic fossil organisms. In some cases, it has proven difficult to assign a problematic fossil even to the invertebrates or vertebrates (more generally chordates) based on often incompletely preserved morphology alone, and chemical composition may help to resolve such questions. Here, we use in situ Raman microspectroscopy to investigate the chemistry of a diverse array of invertebrate and vertebrate fossils from the Pennsylvanian Mazon Creek Lagerstätte of Illinois, and we generate a ChemoSpace through principal component analysis (PCA) of the in situ Raman spectra. Invertebrate soft tissues characterized by chitin (polysaccharide) fossilization products and vertebrate soft tissues characterized by protein fossilization products plot in completely separate, non‐overlapping regions of the ChemoSpace, demonstrating the utility of certain soft tissue molecular signatures as biomarkers for the original soft tissue composition of fossil organisms. The controversial problematicum Tullimonstrum, known as the Tully Monster, groups with the vertebrates, providing strong evidence of a vertebrate rather than invertebrate affinity.