Speciation Rates Research Articles

USING HIGH-RESOLUTION AMMONITE BIOCHRONOSTRATIGRAPHY TO DATE VOLCANOGENIC DEPOSITS PRESERVED IN MIDDLE JURASSIC CARBONATE PLATFORMS SUCCESSIONS OF THE WESTERNMOST TETHYS (SOUTHEASTERN IBERIAN RANGE, SPAIN)

The Middle Jurassic successions that are currently exposed in eastern Spain (southeastern Iberian Range) were deposited in a system of shallow carbonate platforms, constituting the western margin of the Alpine Ocean, and display a series of successively interbedded volcaniclastic rocks and minor lava flows. The large-scale mapping reveals that Middle Jurassic volcanic rocks crop out along NW‒SE pathways, highlighting the importance of the role played by tectonics in controlling the spatial distribution of volcanic outcrops. The progressive dismemberment of Pangea impelled a geographic provinciality increase and, thus, might have contributed to a growing speciation rate of marine invertebrate organisms, notably ammonites. The present study provides new data constraining in detail the age of the Middle Jurassic volcanic deposits. The biochronostratigraphic calibrations of the sedimentary host rocks encasing the volcanics have been obtained from the study of the fossil content (mainly ammonites) from16 well-exposed stratigraphic sections. The study of the fossils from these sections has enabled the identification of three volcanic episodes during the Murchisonae Zone (Aalenian), the Concavum‒Discites zonal boundary (Aalenian-Bajocian), and the late Laeviuscula Zone or Laeviuscula‒Propinquans zonal boundary (Bajocian). Precise age constraints for volcanic accumulations can be a significant contribution in reconstructing the geodynamic history of the Iberian Basin.

Food Plant Shifts Drive the Diversification of Sack-Bearer Moths.

AbstractLepidoptera are a highly diverse group of herbivorous insects; however, some superfamilies have relatively few species. Two alternative hypotheses for drivers of Lepidoptera diversity are shifts in food plant use or shifts from concealed to external feeding as larvae. Many studies address the former hypothesis but with bias toward externally feeding taxa. One of the most striking examples of species disparity between sister lineages in Lepidoptera is between the concealed-feeding sack-bearer moths (Mimallonoidea), which contain about 300 species, and externally feeding Macroheterocera, which have over 74,000 species. We provide the first dated tree of Mimallonidae to understand the diversification dynamics of these moths in order to fill a knowledge gap pertaining to drivers of diversity within an important concealed-feeding clade. We find that Mimallonidae is an ancient Lepidoptera lineage that originated in the Cretaceous ∼105 million years ago and has had a close association with the plant order Myrtales for the past 40 million years. Diversification dynamics are tightly linked with food plant usage in this group. Reliance on Myrtales may have influenced diversification of Mimallonidae because clades that shifted away from the ancestral condition of feeding on Myrtales have the highest speciation rates in the family.

Punctuational ecological changes rather than global factors drive species diversification and the evolution of wing phenotypes inMorphobutterflies

Assessing the relative importance of geographical and ecological drivers of evolution is paramount to understand the diversification of species and traits at the macroevolutionary scale. Here, we use an integrative approach, combining phylogenetics, biogeography, ecology and quantified phenotypes to investigate the drivers of both species and phenotypic diversification of the iconic Neotropical butterfly genus Morpho. We generated a time-calibrated phylogeny for all known species and inferred historical biogeography. We fitted models of time-dependent (accounting for rate heterogeneity across the phylogeny) and paleoenvironment-dependent diversification (accounting for global effect on the phylogeny). We used geometric morphometrics to assess variation of wing size and shape across the tree and investigated their dynamics of evolution. We found that the diversification of Morpho is best explained when considering variable diversification rates across the tree, possibly associated with lineages occupying different microhabitat conditions. First, a shift from understory to canopy was characterized by an increased speciation rate partially coupled with an increasing rate of wing shape evolution. Second, the occupation of dense bamboo thickets accompanying a major host-plant shift from dicotyledons towards monocotyledons was associated with a simultaneous diversification rate shift and an evolutionary 'jump' of wing size. Our study points to a diversification pattern driven by punctuational ecological changes instead of a global driver or biogeographic history.

Geographic patterns and climate correlates of the deviation between phylogenetic and taxonomic diversity for angiosperms in China

Taxonomic diversity (TD) of areas within a region is often strongly and positively correlated with phylogenetic diversity (PD) of the areas. However, variation in speciation and extinction rates and the biogeographic histories of lineages can result in significant deviation of PD from TD (i.e., relative PD after correcting for TD; hereafter PDdev). Identifying areas with higher or lower PDdev within a region is important not only to understanding evolutionary processes associated with the causes of biodiversity patterns across the region but also to identifying areas to be protected for different dimensions of biodiversity. Here, we contrast geographic distributions of PD versus TD for angiosperms in China, explore geographic patterns of PDdev, identify areas where PD is much higher (hotspots) or much lower (coldspots) than expected based on TD, and relate the three diversity metrics to climate variables. We found that geographic patterns of TD were highly congruent with those of PD but they were incongruent with those of PDdev. Areas of unusually high PDdev were generally not those of high TD and PD. Biodiversity hotspots identified based on PDdev generally did not overlap with those identified based on TD and PD. We suggest that identifying areas to be protected should consider not only areas of high TD and PD but also areas of unusually high PDdev, because areas of high PDdev likely hold ancient evolutionary histories and uniqueness, and these areas may not overlap with those having high TD and PD.

An early burst in brachiopod evolution corresponding with significant climatic shifts during the Great Ordovician Biodiversification Event.

We employ modified tip-dating methods to date divergence times within the Strophomenoidea, one of the most abundant and species-rich brachiopod clades to radiate during the Great Ordovician Biodiversification Event (GOBE), to determine if significant environmental changes at this time correlate with the diversification of the clade. Models using origination, extinction and sampling rates to estimate prior probabilities of divergence times strongly support both high rates of anatomical change per million years and rapid divergences shortly before the clade first appears in the fossil record. These divergence times indicate much higher rates of cladogenesis than are typical of brachiopods during this interval. The correspondence of high speciation rates and high anatomical disparity suggests punctuated (speciational) change drove the high frequencies of early anatomical change, which in turn suggests increased ecological opportunities rather than shifting developmental constraints account for high rates of anatomical change. The pulse of rapid evolution began coincident with cooling temperatures, the start of major oscillations in sea level and increased levels of atmospheric oxygen. Our results suggest that these factors permitted major geographical and ecological expansion of strophomenoids with intervals of geographical isolation, resulting in elevated speciation rates and corresponding elevated frequencies of punctuated change.

Niche shifts after island colonization spurred adaptive diversification and speciation in a cosmopolitan bird clade.

Islands have long been recognized as key contributors to biodiversity because they facilitate geographic isolation and ecological divergence from mainland ancestors. However, island colonization has traditionally been considered an evolutionary dead-end process, and its consequences for continental biodiversity remain understudied. Here, we use the evolutionary radiation of Columbiformes (i.e. pigeons and doves) to examine if ecological niche shifts on islands shaped biological diversification and community composition on continents. We show that the colonization of islands by continental, terrestrial-foraging lineages led to the exploitation of a new ecological niche (i.e. arboreal foraging). This transition towards arboreal foraging was associated with evolutionary adaptation towards a new morphological optimum. In addition, arboreal-foraging lineages of islands experienced an increase in speciation rates, which was associated with successful range expansions to other islands as well as back colonization of continents. Our results provide empirical evidence that diversification on continents can only be fully understood when studying the diversification processes that took place on islands, challenging the view of islands as mere sinks of evolutionary diversity.

Novel classification and biogeography ofLeptolejeunea(Lejeuneaceae, Marchantiophyta) with implications for the origin and evolution of the Asian evergreen broad‐leaved forests

AbstractAlthough recent molecular phylogenetic analyses have greatly improved our understanding of the classification of the large liverwort family Lejeuneaceae, the frequent incongruencies between morphology‐based taxonomy and molecular phylogeny have hindered our understanding of evolutionary diversification within the group. Here we focus onLeptolejeunea(Spruce) Steph., a pantropical epiphyllous genus in Lejeuneaceae with 40 species. Phylogenetic studies on the genus have been hampered by insufficient taxon sampling, leaving the deep phylogenetic relationships within this group unresolved. We present the most comprehensive phylogenetic analysis of the genus to date with sampling of over 80% of species, including the enigmaticLeptolejeunea spinistipula(Mizut.) X.L.He endemic to Borneo. Based on data from three molecular markers with representatives ofLeptolejeuneaand its allies,Leptolejeuneaappeared to be monophyletic following the exclusion ofL. spinistipulaand its transfer toSoellaR.L.Zhu, L.Shu, Qiong He & Y.M.Wei. A total‐evidence approach was taken to resolve the backbone phylogeny ofLeptolejeuneaand a first infrageneric classification ofLeptolejeunea, including a new subgenus and three new sections, is proposed based on integrated molecular and morphological evidence. Reconstruction of the evolutionary history showed a wide ancestral area ofLeptolejeuneaduring the Paleogene that arose in mainland Asia, followed by an accelerated speciation rate. Across the biogeographical history ofLeptolejeunea, long‐distance dispersal had profound effects on population expansion. Our findings suggest that Australasia is a source of biodiversity of Asian evergreen broad‐leaved forests that have been established since the Oligocene and rose after the early Miocene.

Human reference gut microbiome catalog including newly assembled genomes from under-represented Asian metagenomes

BackgroundMetagenome sampling bias for geographical location and lifestyle is partially responsible for the incomplete catalog of reference genomes of gut microbial species. Thus, genome assembly from currently under-represented populations may effectively expand the reference gut microbiome and improve taxonomic and functional profiling.MethodsWe assembled genomes using public whole-metagenomic shotgun sequencing (WMS) data for 110 and 645 fecal samples from India and Japan, respectively. In addition, we assembled genomes from newly generated WMS data for 90 fecal samples collected from Korea. Expecting genome assembly for low-abundance species may require a much deeper sequencing than that usually employed, so we performed ultra-deep WMS (> 30 Gbp or > 100 million read pairs) for the fecal samples from Korea. We consequently assembled 29,082 prokaryotic genomes from 845 fecal metagenomes for the three under-represented Asian countries and combined them with the Unified Human Gastrointestinal Genome (UHGG) to generate an expanded catalog, the Human Reference Gut Microbiome (HRGM).ResultsHRGM contains 232,098 non-redundant genomes for 5414 representative prokaryotic species including 780 that are novel, > 103 million unique proteins, and > 274 million single-nucleotide variants. This is an over 10% increase from the UHGG. The new 780 species were enriched for the Bacteroidaceae family, including species associated with high-fiber and seaweed-rich diets. Single-nucleotide variant density was positively associated with the speciation rate of gut commensals. We found that ultra-deep sequencing facilitated the assembly of genomes for low-abundance taxa, and deep sequencing (e.g., > 20 million read pairs) may be needed for the profiling of low-abundance taxa. Importantly, the HRGM significantly improved the taxonomic and functional classification of sequencing reads from fecal samples. Finally, analysis of human self-antigen homologs on the HRGM species genomes suggested that bacterial taxa with high cross-reactivity potential may contribute more to the pathogenesis of gut microbiome-associated diseases than those with low cross-reactivity potential by promoting inflammatory condition.ConclusionsBy including gut metagenomes from previously under-represented Asian countries, Korea, India, and Japan, we developed a substantially expanded microbiome catalog, HRGM. Information of the microbial genomes and coding genes is publicly available (www.mbiomenet.org/HRGM/). HRGM will facilitate the identification and functional analysis of disease-associated gut microbiota.

Mitogenomes of Accipitriformes and Cathartiformes Were Subjected to Ancestral and Recent Duplications Followed by Gradual Degeneration.

The rearrangement of 37 genes with one control region, firstly identified in Gallus gallus mitogenome, is believed to be ancestral for all Aves. However, mitogenomic sequences obtained in recent years revealed that many avian mitogenomes contain duplicated regions that were omitted in previous genomic versions. Their evolution and mechanism of duplication are still poorly understood. The order of Accipitriformes is especially interesting in this context because its representatives contain a duplicated control region in various stages of degeneration. Therefore, we applied an appropriate PCR strategy to look for duplications within the mitogenomes of the early diverged species Sagittarius serpentarius and Cathartiformes, which is a sister order to Accipitriformes. The analyses revealed the same duplicated gene order in all examined taxa and the common ancestor of these groups. The duplicated regions were subjected to gradual degeneration and homogenization during concerted evolution. The latter process occurred recently in the species of Cathartiformes as well as in the early diverged lineages of Accipitriformes, that is, Sagittarius serpentarius and Pandion haliaetus. However, in other lineages, that is, Pernis ptilorhynchus, as well as representatives of Aegypiinae, Aquilinae, and five related subfamilies of Accipitriformes (Accipitrinae, Circinae, Buteoninae, Haliaeetinae, and Milvinae), the duplications were evolving independently for at least 14–47 Myr. Different portions of control regions in Cathartiformes showed conflicting phylogenetic signals indicating that some sections of these regions were homogenized at a frequency higher than the rate of speciation, whereas others have still evolved separately.

Historical warming consistently decreased size, dispersal and speciation rate of fish

There is ongoing debate as to whether fish body size will decrease with global warming and how these changes may impact dispersal ability and speciation rate. Theory predicts that, under warmer temperatures, fish grow to a smaller size, undergo a reduction in dispersal ability and increase speciation rates. However, evaluations of such predictions are hampered owing to the lack of empirical data spanning both wide temporal and geographical scales. Here, using phylogenetic methods, we show that smaller clupeiform fish (anchovies and herrings) occurred historically in warmer waters, moved the shortest distances at low speed and displayed the lowest speciation rates. Furthermore, fish moved faster and evolved rapidly under higher rates of temperature change but these historical rates are far lower than current warming rates. Our results predict a future where smaller clupeiform fish that have reduced ability to move will be more prevalent; this, in turn, may reduce future speciation.

Molecules and fossils tell distinct yet complementary stories of mammal diversification

Reconstructing the tempo at which biodiversity arose is a fundamental goal of evolutionary biologists, yet the relative merits of evolutionary-rate estimates are debated based on whether they are derived from the fossil record or time-calibrated phylogenies (timetrees) of living species. Extinct lineages unsampled in timetrees are known to "pull" speciation rates downward, but the temporal scale at which this bias matters is unclear. To investigate this problem, we compare mammalian diversification-rate signatures in a credible set of molecular timetrees (n= 5,911 species, ∼70% from DNA) to those in fossil genus durations (n= 5,320). We use fossil extinction rates to correct or "push" the timetree-based (pulled) speciation-rate estimates, finding a surge of speciation during the Paleocene (∼66-56 million years ago, Ma) between the Cretaceous-Paleogene (K-Pg) boundary and the Paleocene-Eocene Thermal Maximum (PETM). However, about two-thirds of the K-Pg-to-PETM originating taxa did not leave modern descendants, indicating that this rate signature is likely undetectable from extant lineages alone. For groups without substantial fossil records, thankfully all is not lost. Pushed and pulled speciation rates converge starting ∼10 Ma and are equal at the present day when recent evolutionary processes can be estimated without bias using species-specific "tip" rates of speciation. Clade-wide moments of tip rates also enable enriched inference, as the skewness of tip rates is shown to approximate a clade's extent of past diversification-rate shifts. Molecular timetrees need fossil-correction to address deep-time questions, but they are sufficient for shallower time questions where extinctions are fewer.

Evolutionary dynamics of the elevational diversity gradient in passerine birds.

Low-elevation regions harbour the majority of the world's species diversity compared to high-elevation areas. This global gradient suggests that lowland species have had more time to diversify, or that net diversification rates have been higher in the lowlands. However, highlands seem to be cradles of diversity as they contain many young endemics, suggesting that their rates of speciation are exceptionally fast. Here we use a phylogenetic diversification model that accounts for the dispersal of species between different elevations to examine the evolutionary dynamics of the elevational diversity gradient in passerine birds, a group that has radiated globally to occupy almost all elevations and latitudes. We find strong support for a model in which passerines diversify at the same rate in the highlands and the lowlands but in which the per-capita rate of dispersal from high to low elevations is more than twice as fast as that in the reverse direction. This suggests that while there is no consistent trend in diversification across elevations, part of the diversity generated by highland regions migrates into the lowlands, thus setting up the observed gradient in passerine diversity. We find that this process drives tropical regions but for temperate areas, the analysis could be hampered by their lower richness. Despite their lower diversity, highland regions are disproportionally important for maintaining diversity in the adjacent lowlands.

Phylogenetic analysis and trait evolution of ant cocoons

Abstract Most ant species have lost the ability to spin cocoons. To explore the evolution of cocoon loss within Formicidae, we perform an ancestral state reconstruction of cocooned pupae across a genus-level phylogeny and use a sister clade analysis to determine the impact of cocoon evolution on ant speciation. Then, we fit models of correlated evolution between cocoon status and several other organismal traits. We find that the re-emergence of cocoons is rare and that “naked” lineages display an increased rate of speciation in 5 out of 9 sister group comparisons. Models of correlated evolution with cocoon status were favored for metapleural gland and worker polymorphism. Metapleural gland favored rates of evolution were inconclusive, while worker polymorphism displayed a higher transition rate towards polymorphism coupled with cocoon loss. These results suggest that cocoon loss may allow for other complex traits to develop and may represent a novel example of relaxed selection.

Amphibian Speciation Rates Support a General Role of Mountains as Biodiversity Pumps.

AbstractContinental mountain areas cover <15% of global land surface, yet these regions concentrate >80% of global terrestrial diversity. One prominent hypothesis to explain this pattern proposes that high mountain diversities could be explained by higher diversification rates in regions of high topographic complexity (HTC). While high speciation in mountains has been detected for particular clades and regions, the global extent to which lineages experience faster speciation in mountains remains unknown. Here we addressed this issue using amphibians as a model system (>7,000 species), and we found that families showing high speciation rates contain a high proportion of species distributed in mountains. Moreover, we found that lineages inhabiting areas of HTC speciate faster than lineages occupying areas that are topographically less complex. When comparing across regions, we identified the same pattern in five biogeographical realms where higher speciation rates are associated with higher levels of complex topography. Low-magnitude differences in speciation rates between some low and high complex topographies suggest that high mountain diversity is also affected by low extinction and/or high colonization rates. Nevertheless, our results bolster the importance of mountains as engines of speciation at different geographical scales and highlight their importance for the conservation of global biodiversity.

Comparing diversification rates in lakes, rivers, and the sea.

The diversity of species inhabiting freshwater relative to marine habitats is striking, given that freshwater habitats encompass <1% of Earth's water. The most commonly proposed explanation for this pattern is that freshwater habitats are more fragmented than marine habitats, allowing more opportunities for allopatric speciation and thus increased diversification rates in freshwater. However, speciation may be generally faster in sympatry than in allopatry, as illustrated by lacustrine radiations such as African cichlids. Such differences between rivers and lakes may be important to consider when comparing diversification broadly among freshwater and marine groups. Here I compared diversification rates of teleost fishes in marine, riverine and lacustrine habitats. I found that lakes had faster speciation and net diversification rates than other aquatic habitats. However, most freshwater diversity arose in rivers. Surprisingly, riverine and marine habitats had similar rates of net diversification on average. Biogeographic models suggest that lacustrine habitats are evolutionarily unstable, explaining the dearth of lacustrine species in spite of their rapid diversification. Collectively, these results suggest that strong diversification rate differences are unlikely to explain the freshwater paradox. Instead, this pattern may be attributable to the comparable amount of time spent in riverine and marine habitats over the 200-million-year history of teleosts.

Effect of Productivity on Community Size Explains the Latitudinal Diversity Gradient of South American Small Mammals.

AbstractAlthough many studies have shown that species richness increases from high to low latitudes (the latitudinal diversity gradient), the mechanisms responsible for generating and maintaining higher species richness in the tropics remain intensely debated. Here we investigate how the effects of temperature on speciation rates (kinetic effects) and the effects of productivity on community size (chemical effects) explain the latitudinal diversity gradient of South American small mammals. We implemented Bayesian models that integrate processes from the neutral and metabolic theories, comparing model predictions with empirical richness patterns. The neutral-metabolic model predicted the latitudinal richness gradient in South American small mammals. We found evidence that the effects of productivity on community size are more important for explaining differences in species richness than the effects of temperature on speciation rates. These results suggest that differences in species richness along latitudinal gradients are regulated primarily by the chemical effects of productivity on speciation-extinction dynamics.

A New Late Permian Burnetiamorph From Zambia Confirms Exceptional Levels of Endemism in Burnetiamorpha (Therapsida: Biarmosuchia) and an Updated Paleoenvironmental Interpretation of the Upper Madumabisa Mudstone Formation

A new burnetiamorph therapsid, Isengops luangwensis, gen. et sp. nov., is described on the basis of a partial skull from the upper Madumabisa Mudstone Formation of the Luangwa Basin of northeastern Zambia. Isengops is diagnosed by reduced palatal dentition, a ridge-like palatine-pterygoid boss, a palatal exposure of the jugal that extends far anteriorly, a tall trigonal pyramid-shaped supraorbital boss, and a recess along the dorsal margin of the lateral temporal fenestra. The upper Madumabisa Mudstone Formation was deposited in a rift basin with lithofacies characterized by unchannelized flow, periods of subaerial desiccation and non-deposition, and pedogenesis, and can be biostratigraphically tied to the upper Cistecephalus Assemblage Zone of South Africa, suggesting a Wuchiapingian age. Isengops is the second burnetiamorph recognized from Zambia and is part of a tetrapod assemblage remarkably similar to others across southern Pangea during the Wuchiapingian. A revised cladistic analysis of Biarmosuchia yielded over 500 most parsimonious trees that generally reaffirm the results of previous analyses for burnetiamorphs: Lemurosaurus is basal, Lobalopex and Isengops are proximate burnetiid outgroups, and Bullacephalus, Burnetia, Mobaceras, Niuksenitia, and Pachydectes are burnetiines. Furthermore, Russian biarmosuchians are scattered throughout the tree and do not form sister taxon relationships with each other. Burnetiamorphs display a wide disparity of cranial adornments and are relatively speciose (13 species), especially when compared to the number of specimens discovered to date (∼16 specimens). As has been suggested in some other tetrapod clades (e.g., ceratopsian dinosaurs), the burnetiamorph fossil record supports an inferred macroevolutionary relationship between cranial adornment and increased speciation rate.

The diversification of termites: Inferences from a complete species‐level phylogeny

AbstractTermites play a major role in a variety of ecological processes in tropical and subtropical biomes worldwide, such as decomposition, soil formation and aeration, and nutrient cycling. These important ecosystem services were achieved through their highly complex societies and remarkable adaptations, including the evolution of reproductive division of labour, the acquisition of endosymbionts and the capacity for extensive environmental engineering, yet the causes and consequences of their ecological success are still poorly understood. The goals of our study were (a) to provide the first complete, species‐level phylogeny of all currently recognized termite species by integrating the available genetic and taxonomic data, as well as methods of phylogenetic imputation and divergence time estimation; and (b) to explore variation in speciation rates among termite lineages. We provide the inferred relationships as a set of 1,000 pseudo‐posterior trees, which can be used in future comparative analyses. We demonstrate that speciation rates have been relatively constant throughout the history of termites, with two positive shifts in speciation rates: one at their origin of Euisoptera and the other concordant with evolution of Termitidae. On the other hand, there was no obvious trend towards deceleration in speciation rates for termites as a whole, nor within the most species‐rich families. The provided trees might represent a valuable resource for termite comparative studies by summarizing the available phylogenetic information, while accounting for uncertainty in the inferred topologies.

Planktonic foraminiferal biostratigraphy of the Cretaceous strata, Indus Basin, Pakistan, Eastern Tethys: implications for oceanic anoxic events

Stratigraphic horizons of the Cretaceous oceanic anoxic events (OAEs) are constrained for the first time at the northwestern margin of the Indian Plate, Pakistan, using high-resolution planktonic foraminiferal biostratigraphy. In this study, 15 planktonic foraminiferal biozones have been erected, suggesting an early Aptian to early Maastrichtian age for the studied section. Based on the established biozones, the stratigraphic positions of the Cretaceous OAEs are defined. The lowermost organic-rich interval occurs in the Hedbergella delrioensis and Globigerinelloides algerianus zones of the early to late Aptian, which correlate with the Selli (OAE 1a) and Fallot events. The next interval with the abundant presence of organic matter occurs in the Ticinella bejaouaensis zone (latest Aptian) and correlates with OAE1b. The third organic-rich interval falls in the Biticinella breggiensis zone of the middle to late Albian and correlates with the OAE1c. The fourth organic-rich horizon straddles the Cenomanian − Turonian boundary and corresponds to OAE2. The fifth organic-rich interval falls in the Globotruncana ventricosa zone of middle to late Campanian age and correlates with the Campanian isotopic event. The evolution of planktonic foraminifera in this part of the Tethys Ocean, is driven by OAEs, e.g. at the base of OAE2, a mass extinction of foraminiferal species, mainly rotaliporids, occurred, and at the end of OAE2, simultaneous appearances of the whiteinellids, marginotruncanids and dicarinellids indicate the fastest rate of speciation. KEY POINTS The planktonic foraminiferal biostratigraphic investigations revealed an early Aptian to early Maastrichtian age for the Parh Formation exposed in the Mughal Kot Section, Pakistan. Based on biostratigraphic data, stratigraphic positions of the OAE1a, OAE1b, OAE1c, OAE2 and Campanian events are constrained. An evolutionary link of major planktonic foraminiferal genera with Cretaceous OAEs has been observed.

Diversification dynamics of total-, stem-, and crown-groups are compatible with molecular clock estimates of divergence times.

Molecular evolutionary time scales are expected to predate the fossil evidence, but, particularly for major evolutionary radiations, they can imply extremely protracted stem lineages predating the origin of living clades, leading to claims of systematic overestimation of divergence times. We use macroevolutionary birth-death models to describe the range of total-group and crown-group ages expected under constant rates of speciation and extinction. We extend current predictions on origination times for crown- and total-groups, and extinction of stem-groups, demonstrating that there is broad variance in these predictions. Under constant rates of speciation and extinction, we show that the distribution of expected arthropod total-group ages is consistent with molecular clock estimates. The fossil record cannot be read literally, and our results preclude attempts to interpret the antiquity of clades based on the co-occurrence of stem- and crown-representatives.