Extinction and time help drive the marine‐terrestrial biodiversity gradient: is the ocean a deathtrap?

The marine‐terrestrial richness gradient is among Earth's most dramatic biodiversity patterns, but its causes remain poorly understood. Here, we analyse detailed phylogenies of amniote clades, paleontological data and simulations to reveal the mechanisms underlying low marine richness, emphasising speciation, extinction and colonisation. We show that differences in diversification rates (speciation minus extinction) between habitats are often weak and inconsistent with observed richness patterns. Instead, the richness gradient is explained by limited time for speciation in marine habitats, since all extant marine clades are relatively young. Paleontological data show that older marine invasions have consistently ended in extinction. Simulations show that marine extinctions help drive the pattern of young, depauperate marine clades. This role for extinction is not discernible from molecular phylogenies alone, and not predicted by most previously hypothesised explanations for this gradient. Our results have important implications for the marine‐terrestrial biodiversity gradient, and studies of biodiversity gradients in general.     

Marine planktonic microbes survived climatic instabilities in the past

In the geological past, changes in climate and tectonic activity are thought to have spurred the tempo of evolutionary change among major taxonomic groups of plants and animals. However, the extent to which these historical contingencies increased the risk of extinction of microbial plankton species remains largely unknown. Here, I analyse fossil records of marine planktonic diatoms and calcareous nannoplankton over the past 65 million years from the world oceans and show that the probability of species' extinction is not correlated with secular changes in climatic instability. Further supporting these results, analyses of genera survivorship curves based on fossil data concurred with the predictions of a birth-death model that simulates the extinction of genera through time assuming stochastically constant rates of speciation and extinction. However, my results also show that these marine microbes responded to exceptional climatic contingencies in a manner that appears to have promoted net diversification. These results highlight the ability of marine planktonic microbes to survive climatic instabilities in the geological past, and point to different mechanisms underlying the processes of speciation and extinction in these micro-organisms.     

Terrestrial nemerteans thirty years on

The present paper records changes in the reported distribution and abundance of terrestrial nemerteans during the last 30 years. New records are reported from the Isles of Scilly, Bermuda and New Zealand of species previously only known elsewhere. Human activity, especially the increased importation and exportation of plants between countries, has probably led to terrestrial nemerteans, as well as other organisms, becoming introduced into many new localities. However, the conservation outlook is not hopeful because of a global loss of appropriate habitat, particularly forests. Today three species (Geonemertes rodericana Gulliver, 1879, Pantinonemertes agricola Willemoes-Suhm, 1874 and Antiponemertes allisonae Moore, 1973) are feared to be extinct, and none of the extant species seem to be as abundant as they once were. Behavioural changes in Antiponemertes pantini (Southgate, 1954) under arid conditions may be associated with heavy gregarine infestations.     

Molecular phylogeny of Enchytraeidae (Oligochaeta) indicates separate invasions of the terrestrial environment

Enchytraeidae is a family of soil inhabiting small‐ to medium‐sized oligochaete worms using degradable plant material as a food source and primarily adapted to terrestrial or semi‐terrestrial environments. The molecular phylogeny based upon both mitochondrial and nuclear genes indicates early segregations of the two genera Enchytraeus and Lumbricillus leaving the remaining genera included in this study as a later segregated major monophyletic branch. Extant members of the two former genera dominate in decaying seaweed in the littoral zone along the sea although members of in particular the genus Enchytraeus have also invaded other habitats. Historically the littoral zone of the sea is undoubtedly the first terrestrial or semi‐terrestrial habitat where dead plant material accumulates to any greater extent and Enchytraeus and Lumbricillus may represent early successful attempts to exploit this resource. Inland soils probably had to await the emergence of land plants in order to provide a similar food resource and here the major branch of enchytraeid genera diversified into a high number of species in the numerous decomposer networks of this varied environment. A subdivision into the genera Enchytraeus and Lumbricillus on the one hand and a branch of mainly inland genera on the other is supported by differences in two somewhat neglected morphological features. Firstly, in Enchytraeus and Lumbricillus the testes are enclosed in a testis sac within which the male cells mature, by one possible exception a unique feature among Oligochaeta, The other enchytraeid genera studied and Oligochaeta in general lack this sac and the male cells mature directly in the cavity of the testicular segment. Secondly, species of Enchytraeus and Lumbricillus generally have a higher reproductive output than species of the inland terrestrial branch and this may represent an adaptation to the unpredictable littoral zone compared to the more stable nature of inland habitats. In the older literature the genus Mesenchytraeus is considered to have a basic position within the entire family but our molecular data do not support this expectation. In Enchytraeidae the nephridia are elaborate organs of a characteristic and constant shape covering species from different genera in a pattern following the molecular phylogeny. Other much used morphological features such as shape of setae, anteclitellar origin of the dorsal vessel and various modifications of the intestine have arisen more than once.     

Detecting shifts in diversity limits from molecular phylogenies: what can we know?

Large complete species-level molecular phylogenies can provide the most direct information about the macroevolutionary history of clades having poor fossil records. However, extinction will ultimately erode evidence of pulses of rapid speciation in the deep past. Assessment of how well, and for how long, phylogenies retain the signature of such pulses has hitherto been based on a--probably untenable--model of ongoing diversity-independent diversification. Here, we develop two new tests for changes in diversification 'rules' and evaluate their power to detect sudden increases in equilibrium diversity in clades simulated with diversity-dependent speciation and extinction rates. Pulses of diversification are only detected easily if they occurred recently and if the rate of species turnover at equilibrium is low; rates reported for fossil mammals suggest that the power to detect a doubling of species diversity falls to 50 per cent after less than 50 Myr even with a perfect phylogeny of extant species. Extinction does eventually draw a veil over past dynamics, suggesting that some questions are beyond the limits of inference, but sudden clade-wide pulses of speciation can be detected after many millions of years, even when overall diversity is constrained. Applying our methods to existing phylogenies of mammals and angiosperms identifies intervals of elevated diversification in each.     

To what extent do new fossil discoveries change our understanding of clade evolution? A cautionary tale from burying beetles (Coleoptera: Nicrophorus)

Divergence time estimates derived from phylogenies are crucial to infer historical biogeography and diversification dynamics. Yet, the impact of fossil record incompleteness on macroevolutionary reconstructions remains equivocal. Here, we investigate to what extent gaps in the fossil record can impinge downstream evolutionary inferences in the beetle family Silphidae. Recent discoveries have pushed back the fossil record of this group from the Eocene into the Jurassic. We estimated the divergence times of the family using both its currently understood fossil record and the fossil record known prior to these recent discoveries. All fossil calibrations were informed with different parametric distributions to investigate the weight of priors on posterior age estimates. Based on time‐calibrated trees, we assessed the impact of fossil calibrations on the inference of ancestral ranges and diversification rate dynamics in the genus Nicrophorus. Depending upon the selected sets of fossil constraints, the age discrepancies had a major impact on the macroevolutionary inferences: the biogeographic extrapolations relative to paleogeography are markedly contrasting, and the calculated rates at which species form or go extinct (and when they varied) are strikingly different. We show that soft prior distributions do not necessarily alleviate such shortcomings therefore preventing the inference of reliable macroevolutionary patterns in groups presenting a taphonomic bias in their fossil record.     

Nymphalid butterflies diversify following near demise at the Cretaceous/Tertiary boundary

The butterfly family Nymphalidae contains some of the most important non-drosophilid insect model systems for evolutionary and ecological studies, yet the evolutionary history of the group has remained shrouded in mystery. We have inferred a robust phylogenetic hypothesis based on sequences of 10 genes and 235 morphological characters for exemplars of 400 of the 540 valid nymphalid genera representing all major lineages of the family. By dating the branching events, we infer that Nymphalidae originated in the Cretaceous at 90 Ma, but that the ancestors of 10–12 lineages survived the end-Cretaceous catastrophe in the Neotropical and Oriental regions. Patterns of diversification suggest extinction of lineages at the Cretaceous/Tertiary boundary (65 Ma) and subsequent elevated speciation rates in the Tertiary.     

Extinction and diversification in the Devonian Brachiopoda of New York state: No correlation with sea level?

Sea level highstand is generally considered to promote high species diversities among marine organisms through habitat expansion and global climatic amelioration, and marine regression to trigger elevated extinction rates among marine benthic organisms by habitat reduction (the species‐area effect), and among both marine and terrestrial organisms by global climatic deterioration. The Devonian is unusual in that the Late Devonian mass extinction occurs during an interval of global sea level highstand. To further explore this anomaly, the potential relationship between relative sea level and evolutionary biology is analyzed here for the Brachiopoda of the Devonian Period. Successive linear modeling reveals a total lack of correlation between relative sea level and either origination rates, extinction rates, or standing diversity among the Devonian brachiopods.     

Phylogenetic diversity as a window into the evolutionary and biogeographic histories of present-day richness gradients for mammals

Phylogenetic diversity (PD) captures the shared ancestry of species, and is increasingly being recognized as a valuable conservation currency. Regionally, PD frequently covaries closely with species richness; however, variation in speciation and extinction rates and/or the biogeographic history of lineages can result in significant deviation. Locally, these differences may be pronounced. Rapid recent speciation or high temporal turnover of lineages can result in low PD but high richness. In contrast, rare dispersal events, for example, between biomes, can elevate PD but have only small impact on richness. To date, environmental predictors of species richness have been well studied but global models explaining variation in PD are lacking. Here, we contrast the global distribution of PD versus species richness for terrestrial mammals. We show that an environmental model of lineage diversification can predict well the discrepancy in the distribution of these two variables in some places, for example, South America and Africa but not others, such as Southeast Asia. When we have information on multiple diversity indices, conservation efforts directed towards maximizing one currency or another (e.g. species richness versus PD) should also consider the underlying processes that have shaped their distributions.