Amazonian Amphibian Diversity Is Primarily Derived from Late Miocene Andean Lineages

The Neotropics contains half of remaining rainforests and Earth's largest reservoir of amphibian biodiversity. However, determinants of Neotropical biodiversity (i.e., vicariance, dispersals, extinctions, and radiations) earlier than the Quaternary are largely unstudied. Using a novel method of ancestral area reconstruction and relaxed Bayesian clock analyses, we reconstructed the biogeography of the poison frog clade (Dendrobatidae). We rejected an Amazonian center-of-origin in favor of a complex connectivity model expanding over the Neotropics. We inferred 14 dispersals into and 18 out of Amazonia to adjacent regions; the Andes were the major source of dispersals into Amazonia. We found three episodes of lineage dispersal with two interleaved periods of vicariant events between South and Central America. During the late Miocene, Amazonian, and Central American-Chocoan lineages significantly increased their diversity compared to the Andean and Guianan-Venezuelan-Brazilian Shield counterparts. Significant percentage of dendrobatid diversity in Amazonia and Chocó resulted from repeated immigrations, with radiations at <10.0 million years ago (MYA), rather than in situ diversification. In contrast, the Andes, Venezuelan Highlands, and Guiana Shield have undergone extended in situ diversification at near constant rate since the Oligocene. The effects of Miocene paleogeographic events on Neotropical diversification dynamics provided the framework under which Quaternary patterns of endemism evolved.     

The complex effects of mass extinctions on morphological disparity

Studies of biodiversity through deep time have been a staple for biologists and paleontologists for over 60 years. Investigations of species richness (diversity) revealed that at least five mass extinctions punctuated the last half billion years, each seeing the rapid demise of a large proportion of contemporary taxa. In contrast to diversity, the response of morphological diversity (disparity) to mass extinctions is unclear. Generally, diversity and disparity are decoupled, such that diversity may decline as morphological disparity increases, and vice versa. Here, we develop simulations to model disparity changes across mass extinctions using continuous traits and birth-death trees. We find no simple null for disparity change following a mass extinction but do observe general patterns. The range of trait values decreases following either random or trait-selective mass extinctions, whereas variance and the density of morphospace occupation only decline following trait-selective events. General trends may differentiate random and trait-selective mass extinctions, but methods struggle to identify trait selectivity. Long-term effects of mass extinction trait selectivity change support for phylogenetic comparative methods away from the simulated Brownian motion toward Ornstein-Uhlenbeck and Early Burst models. We find that morphological change over mass extinction is best studied by quantifying multiple aspects of morphospace occupation.     

Trends and patterns in the evolution of vascular plants: macroevolutionary implications of a multilevel taxonomic analysis

Cascales‐Miñana, B., Muñoz‐Bertomeu, J., Ros, R., Segura, J. 2010: Trends and patterns in the evolution of vascular plants: macroevolutionary implications of a multilevel taxonomic analysis. Lethaia, 10.1111/j.1502‐3931.2009.00212.x Studying the macroevolutionary patterns of vascular plants from the Silurian to the present‐day provides a global record of plant life history. Evolutionary rates (origination, extinction and diversification) for families, orders, classes and divisions were analysed, as was abundance and richness for 21 time intervals. An accumulative analysis, based on the total plant fossil record, the accumulated extinctions and relative diversity, was also carried out. The diversification rate shows a uniquely constant and progressive reduction from the end of the Carboniferous to the Permian when the lowest values are registered. Very small peaks seem to reflect Cretaceous extinction for families. At family level, only two time intervals present higher extinctions, than originations. Richness and accumulative analyses reveal that only 32% of the families analysed became extinct, and that approximately 90% of them disappeared at the end of the Palaeozoic. Our results indicate that plants did not undergo mass extinction events in the ‘big five’ sense, but rather, mass ecological reorganization the absence of important extinction events or evolutionary innovations producing diversification patterns without abrupt changes. □Diversification, evolutionary, extinction, fossil record, innovations, radiation, vascular plants.     

Biotic and abiotic factors driving the diversification dynamics of Crocodylia

Species diversity patterns are governed by complex interactions among biotic and abiotic factors over time and space, but are essentially the result of the diversification dynamics (differential speciation and extinction rates) over the long-term evolutionary history of a clade. Previous studies have suggested that temporal variation in global temperature drove long-term diversity changes in Crocodylia, a monophyletic group of large ectothermic organisms. We use a large database of crocodylian fossil occurrences (192 spp.) and body mass estimations, under a taxic approach, to characterize the global diversification dynamics of crocodylians since the Cretaceous, and their correlation with multiple biotic and abiotic factors in a Bayesian framework. The diversification dynamic of crocodylians, which appears to have originated in the Turonian (c. 92.5 Ma), is characterized by several phases with high extinction and speciation rates within a predominantly low long-term mean rate. Our results reveal long-term diversification dynamics of Crocodylia to be a highly complex process driven by a combination of biotic and abiotic factors which influenced the speciation and extinction rates in dissimilar ways. Higher crocodylian extinction rates are related to low body mass disparity, indicating selective extinctions of taxa at both ends of the body mass spectrum. Speciation rate slowdowns are noted when the diversity of the clade is high and the warm temperate climatic belt is reduced. Our finding supports the idea that temporal variations of body mass disparity, self-diversity, and the warm climate belt size provided more direct mechanistic explanations for crocodylian diversification than do proxies of global temperature.     

Extinction patterns in the avifauna of the Hawaiian islands

Through the continuing accumulation of fossil evidence, it is clear that the avifauna of the Hawaiian Islands underwent a large‐scale extinction event around the time of Polynesian arrival. A second wave of extinctions since European colonization has further altered this unique avifauna. Here I present the first systematic analysis of the factors characterizing the species that went extinct in each time period and those that survived in order to provide a clearer picture of the possible causal mechanisms. These analyses were based on mean body size, dietary and ecological information and phylogenetic lineage of all known indigenous, non‐migratory land and freshwater bird species of the five largest Hawaiian Islands. Extinct species were divided into ‘prehistoric’ and ‘historic’ extinction categories based on the timing of their last occurrence. A model of fossil preservation bias was also incorporated. I used regression trees to predict probability of prehistoric and historic extinction based on ecological variables. Prehistoric extinctions showed a strong bias toward larger body sizes and flightless, ground‐nesting species, even after accounting for preservation bias. Many small, specialized species, mostly granivores and frugivores, also disappeared, implicating a wide suite of human impacts including destruction of dry forest habitat. In contrast, the highest extinction rates in the historic period were in medium‐sized nectarivorous and insectivorous species. These differences result from different causal mechanisms underlying the two waves of extinction.     

Fossil and phylogenetic analyses reveal recurrent periods of diversification and extinction in dictyopteran insects

Variations of speciation and extinction rates determine the fate of clades through time. Periods of high diversification and extinction (possibly mass-extinction events) can punctuate the evolutionary history of various clades, but they remain loosely defined for many biological groups, especially nonmarine invertebrates like insects. Here, we examine whether the cockroaches, mantises and termites (altogether included in Dictyoptera) have experienced episodic pulses of speciation or extinction and how these pulses may be associated with environmental fluctuations or mass extinctions. We relied on molecular phylogeny and fossil data to shed light on the times and rates at which dictyopterans diversified. The diversification of Dictyoptera has alternated between (i) periods of high diversification in the late Carboniferous, Early–Middle Triassic, Early Cretaceous and middle Palaeogene, and (ii) periods of high extinction rates particularly at the Permian-Triassic boundary, but not necessarily correlated with the major global biodiversity crises as in the mid-Cretaceous. This study advocates the importance of analyzing, when possible, both molecular phylogeny and fossil data to unveil diversification and extinction periods for a given group. The causes and consequences of extinction must be studied beyond mass-extinction events alone to gain a broader understanding of how clades wax and wane.     

HOW DID LIFE BECOME SO DIVERSE? THE DYNAMICS OF DIVERSIFICATION ACCORDING TO THE FOSSIL RECORD AND MOLECULAR PHYLOGENETICS

Abstract: The long‐term diversification of life probably cannot be modelled as a simple equilibrial process: the time scales are too long, the potential for exploring new ecospace is too large and it is unlikely that ecological controls can act at global scales. The sum of many clade expansions and reductions, each of which happens according to its own dynamic, probably approximates more a damped exponential curve when translated into a global‐scale species diversification curve. Unfortunately, it is not possible to plot such a meaningful global‐scale species diversification curve through time, but curves at higher taxonomic levels have been produced. These curves are subject to the vagaries of the fossil record, but it is unlikely that the sources of error entirely overwhelm the biological signal. Clades radiate when the external and internal conditions are right: a new territory or ecospace becomes available, and the lineage has acquired a number of characters that open up a new diet or mode of life. Modern high levels of diversity in certain speciose clades may depend on such ancient opportunities taken. Dramatic climatic changes through the Quaternary must have driven extinctions and originations, but many species responded simply by moving to more favourable locations. Ecological communities appear to be no more than merely chance associations of species, but there may be real interactions among species. Ironically, high species diversity may lead to more speciation, not, as had been assumed, less: more species create more opportunities and selective pressures for other species to respond to, rather than capping diversity at a fixed equilibrium level. Studies from the scale of modern ecosystems to global long‐term patterns in the fossil record support a model for the exponential diversification of life, and one explanation for a pattern of exponential diversification is that as diversity increases, new forms become ever more refinements of existing forms. In a sense the world becomes increasingly divided into finer niche space. Organisms have a propensity to speciate freely, species richness within ecosystems appears to generate opportunities for more speciation, clades show all kinds of patterns from sluggish speciation rates and constant diversity through time to apparently explosive speciation, and there is no evidence that rapidly speciating clades have reached a limit, nor that they are driving other clades to extinction. A corollary of this view is that current biodiversity must be higher than it has ever been. Limits to infinite growth are clearly local, regional, and global turnover and extinction events, when climate change and physical catastrophes knock out species and whole clades, and push the rising exponential curve down a notch or two.     

CLIMATE PREDICTORS OF LATE QUATERNARY EXTINCTIONS

Between 50,000 and 3,000 years before present (BP) 65% of mammal genera weighing over 44 kg went extinct, together with a lower proportion of small mammals. Why species went extinct in such large numbers is hotly debated. One of the arguments proposes that climate changes underlie Late Quaternary extinctions, but global quantitative evidence for this hypothesis is still lacking. We test the potential role of global climate change on the extinction of mammals during the Late Quaternary. Our results suggest that continents with the highest climate footprint values, in other words, with climate changes of greater magnitudes during the Late Quaternary, witnessed more extinctions than continents with lower climate footprint values, with the exception of South America. Our results are consistent across species with different body masses, reinforcing the view that past climate changes contributed to global extinctions. Our model outputs, the climate change footprint dataset, provide a new research venue to test hypotheses about biodiversity dynamics during the Late Quaternary from the genetic to the species richness level.     

Biodiversity is not (and never has been) a bed of roses!

Over the last decades, the critical study of fossil diversity has led to significant advances in the knowledge of global macroevolutionary patterns of biodiversity. The deep-time history of life on Earth results from background originations and extinctions defining a steady-state, nonstationary equilibrium occasionally perturbed by biotic crises and "explosive" diversifications. More recently, a macroecological approach to the large-scale distribution of extant biodiversity offered new, stimulating perspectives on old theoretical questions and current practical problems in conservation biology. However, time and space are practically distinct, but functionally related dimensions of ecological systems. This calls for a spatially-integrated study of biodiversity dynamics at an evolutionary timescale. Indeed, the biosphere is a complex adaptive system whose study cannot be arbitrarily reduced to any single spatial- and/or temporal-scale level of resolution without a loss of content. From such an integrated perspective, a simple fact emerges: in a physically heterogeneous and ever-changing world, spatiotemporal variations in biodiversity are the rule-not the exception.     

Body mass‐related changes in mammal community assembly patterns during the late Quaternary of North America

The late Quaternary of North America was marked by prominent ecological changes, including the end‐Pleistocene megafaunal extinction, the spread of human settlements and the rise of agriculture. Here we examine the mechanistic reasons for temporal changes in mammal species association and body size during this time period. Building upon the co‐occurrence results from Lyons et al. (2016) – wherein each species pair was classified as spatially aggregated, segregated or random – we examined body mass differences (BMD) between each species pair for each association type and time period (Late Pleistocene: 40 000 ¹⁴C–11 700 ¹⁴C ybp, Holocene: 11 700 ¹⁴C–50 ybp and Modern: 50–0 yr). In the Late Pleistocene and Holocene, the BMD of both aggregated and segregated species pairs was significantly smaller than the BMD of random pairs. These results are consistent with environmental filtering and competition as important drivers of community structure in both time periods. Modern assemblages showed a breakdown between BMD and co‐occurrence patterns: the average BMD of aggregated, segregated and random species pairs did not differ from each other. Collectively, these results indicate that the late Quaternary mammalian extinctions not only eliminated many large‐bodied species but were followed by a re‐organization of communities that altered patterns of species coexistence and associated differences in body size.     

Phylogenetic Clustering of Origination and Extinction across the Late Ordovician Mass Extinction

Mass extinctions can have dramatic effects on the trajectory of life, but in some cases the effects can be relatively small even when extinction rates are high. For example, the Late Ordovician mass extinction is the second most severe in terms of the proportion of genera eliminated, yet is noted for the lack of ecological consequences and shifts in clade dominance. By comparison, the end-Cretaceous mass extinction was less severe but eliminated several major clades while some rare surviving clades diversified in the Paleogene. This disconnect may be better understood by incorporating the phylogenetic relatedness of taxa into studies of mass extinctions, as the factors driving extinction and recovery are thought to be phylogenetically conserved and should therefore promote both origination and extinction of closely related taxa. Here, we test whether there was phylogenetic selectivity in extinction and origination using brachiopod genera from the Middle Ordovician through the Devonian. Using an index of taxonomic clustering (R_CL) as a proxy for phylogenetic clustering, we find that A) both extinctions and originations shift from taxonomically random or weakly clustered within families in the Ordovician to strongly clustered in the Silurian and Devonian, beginning with the recovery following the Late Ordovician mass extinction, and B) the Late Ordovician mass extinction was itself only weakly clustered. Both results stand in stark contrast to Cretaceous-Cenozoic bivalves, which showed significant levels of taxonomic clustering of extinctions in the Cretaceous, including strong clustering in the mass extinction, but taxonomically random extinctions in the Cenozoic. The contrasting patterns between the Late Ordovician and end-Cretaceous events suggest a complex relationship between the phylogenetic selectivity of mass extinctions and the long-term phylogenetic signal in origination and extinction patterns.     

The two Early Toarcian (Early Jurassic) extinction events in ammonoids

The Early Toarcian (Early Jurassic) biological crisis was one of the ‘minor’ mass extinctions. It is linked with an oceanic anoxic event. Fossil data from sections located in northwestern European (epicontinental platforms and basins) and Tethyan (distal, epioceanic) areas indicate that Late Pliensbachian–Early Toarcian ammonoids experienced two extinction events during the Early Toarcian. The older one is linked with disruption of the Tethyan–Boreal provinciality, whereas the younger event correlates with the onset of anoxia and corresponds with the Early Toarcian mass‐extinction event. These two extinctions cannot be interpreted as episodes of a single, stepwise, event. Values of the net diversification, more than the number of extinctions, allow the two extinction events to be clearly recognized and distinguished. Values of regional net diversification for northwestern European and Tethyan faunas point to greater evolutionary dynamics in the epioceanic areas. The inclusion of Mediterranean faunas in the database proves that the ammonite turnover at the Early Toarcian mass‐extinction event was more important than previously thought. Progenitor (evolute Neolioceratoides), survivor (Dactylioceras, Polyplectus pluricostatus) and Lazarus (Procliviceras) taxa have been recognized. Different selectivity patterns are shown for the two events. The first one, linked to the disruption of the Tethyan–Boreal provinciality, has mainly affected ammonites adapted to epicontinental platforms. In the mass‐extinction event, no selectivity is recognized, because also Phylloceratina and Lytoceratina were deeply affected at species level, although their wide biogeographical distribution at clade level was a significant buffer against extinction. In contrast to Palaeozoic mass extinctions, ammonoid survivors and Lazarus taxa are characterized by complex sutures: Phylloceratina (long‐ranging ammonoids) and Polyplectus (relatively long‐ranging compared to other Ammonitina).     

Effects of extinction on food web structures on an evolutionary time scale

Extinction affected food web structure in paleoecosystems. Recent theoretical studies that examined the effects of extinction intensity on food web structure on ecological time scales have considered extinction to involve episodic events, with pre-extinction food webs becoming established without dynamics. However, in terms of the paleontological time scale, food web structures are generated from feedback with repeated extinctions, because extinction frequency is affected by food web structure, and food web structure itself is a product of previous extinctions. We constructed a simulation model of changes in tri-trophic-level food webs to examine how continual extinction events affect food webs on an evolutionary time scale. We showed that under high extinction intensity (1) species diversity, especially that of consumer species, decreased; (2) the total population density at each trophic level decreased, while the densities of individual species increased; and (3) the trophic link density of the food web increased. In contrast to previous models, our results were based on an assumption of long-term food web development and are able to explain overall trends posited by empirical investigations based on fossil records.     

Lotka's Game in Predator–Prey Theory: Linking Populations to Individuals

This paper further examines an individual-based model of a spatially distributed predator–prey population that demonstrates strong spatial structuring in contrast with predictions from its representative analytic formulation. Examination of a small, localized population reveals that extinctions due to demographic stochasticity dominate the dynamics. Local extinction dynamics produce wave pulses and the interactions of these wave pulses constitute global dynamics. The results motivate a population-level cell-based model with each cell representing a local population and parameterized by local extinction probabilities, rather than individual-based interaction rates. A detailed comparison of spatiotemporal plots from the two modelling frameworks shows that the population-level model captures the broad range of dynamics exhibited by the individual-based model. The agreement between these two complementary theoretical frameworks, one formulated at the level of individuals, the other at the level of populations, provides a mechanistic understanding of the dynamics.     

The sea as deathtrap: comment on a paper by miller and wiens

Miller & Wiens (2017) claim that low marine as compared with terrestrial diversity results from more frequent extinctions and insufficient time for diversification in marine clades. Their data on marine amniotes are unrepresentative of marine diversity, their analysis of clade dynamics is flawed, and they ignore previously proposed explanations for the diversity difference.     

EXPLOSIVE RADIATION OR CRYPTIC MASS EXTINCTION? INTERPRETING SIGNATURES IN MOLECULAR PHYLOGENIES

How biodiversity is generated and maintained underlies many major questions in evolutionary biology, particularly relating to the tempo and pattern of diversification through time. Molecular phylogenies and new analytical methods provide additional tools to help interpret evolutionary processes. Evolutionary rates in lineages sometimes appear punctuated, and such "explosive" radiations are commonly interpreted as adaptive, leading to causative key innovations being sought. Here we argue that an alternative process might explain apparently rapid radiations ("broom-and-handle" or "stemmy" patterns seen in many phylogenies) with no need to invoke dramatic increase in the rate of diversification. We use simulations to show that mass extinction events can produce the same phylogenetic pattern as that currently being interpreted as due to an adaptive radiation. By comparing simulated and empirical phylogenies of Australian and southern African legumes, we find evidence for coincident mass extinctions in multiple lineages that could have resulted from global climate change at the end of the Eocene.     

Determinants of loss of mammal species during the Late Quaternary 'megafauna' extinctions: life history and ecology,but not body size

Extinctions of megafauna species during the Late Quaternary dramatically reduced the global diversity of mammals. There is intense debate over the causes of these extinctions, especially regarding the extent to which humans were involved. Most previous analyses of this question have focused on chronologies of extinction and on the archaeological evidence for human-megafauna interaction. Here, I take an alternative approach: comparison of the biological traits of extinct species with those of survivors. I use this to demonstrate two general features of the selectivity of Late Quaternary mammal extinctions in Australia, Eurasia, the Americas and Madagascar. First, large size was not directly related to risk of extinction; rather, species with slow reproductive rates were at high risk regardless of their body size. This finding rejects the 'blitzkrieg' model of overkill, in which extinctions were completed during brief intervals of selective hunting of large-bodied prey. Second, species that survived despite having low reproductive rates typically occurred in closed habitats and many were arboreal or nocturnal. Such traits would have reduced their exposure to direct interaction with people. Therefore, although this analysis rejects blitzkrieg as a general scenario for the mammal megafauna extinctions, it is consistent with extinctions being due to interaction with human populations.     

Modeling bivalve diversification: the effect of interaction on a macroevolutionary system

The global diversification of the class Bivalvia has historically received two conflicting interpretations. One is that a major upturn in diversification was associated with, and a consequence of, the Lake Permian mass extinction. The other is that mass extinctions have had little influence and that bivalves have experienced slow but nearly steady exponential diversification through most of their history, unaffected by interactions with other clades. We find that the most likely explanation lies between these two interpretations. Through most of the Phanerozoic, the diversity of bivalves did indeed exhibit slow growth, which was not substantially altered by mass extinctions. However, the presence of "hyperexponential bursts" in diversification during the initial Ordovician radiation and following the Late Permian and Late Cretaceous mass extinctions suggests a more complex history in which a higher characteristic diversification rate was dampened through most of the Phanerozoic. The observed pattern can be accounted for with a two-phase coupled (i.e., interactive) logistic model, where one phase is treated as the "bivalves" and the other phase is treated as a hypothetical group of clades with which the "bivalves" might have interacted. Results of this analysis suggest that interactions with other taxa have substantially affected bivalve global diversity through the Phanerozoic.     

Do life history traits account for diversity of polychaete annelids?

Abstract. Within many phylogenetic assemblages, a pattern of domination has been observed: one or a few clades have had many more speciation events or fewer extinctions than other clades in a particular assemblage. We investigated this phenomenon in the polychaete annelids. Polychaetes comprise ∼9000 described species classified in over 70 families and exhibit a great variety of life history strategies. Our goal was to test whether diverse polychaete families are characterized by species with short generation times, high reproductive output, small body size, or with planktotrophic larval development. Each of these factors has been advanced as cause for high diversity in other taxonomic assemblages. Here, we establish that the diversification pattern of polychaete families is non-random, but the data collected show no significant correlations between familial diversity and several life history traits including age at first reproduction, life span, body size, fecundity, and egg size. Pairwise comparisons of sister families do not reveal any trends between familial diversity and any of the life history traits. The great variability of life history traits within polychaete families may explain the lack of significant results; perhaps no trends are seen because polychaete life history traits cannot be generalized at the family level.