Parental care results in a greater mutation load, for which it is also a phenotypic antidote.

Benevolent social behaviours, such as parental care, are thought to enable mildly deleterious mutations to persist. We tested this prediction experimentally using the burying beetle Nicrophorus vespilloides, an insect with biparental care. For 20 generations, we allowed replicate experimental burying beetle populations to evolve either with post-hatching care ('Full Care' populations) or without it ('No Care' populations). We then established new lineages, seeded from these experimental populations, which we inbred to assess their mutation load. Outbred lineages served as controls. We also tested whether the deleterious effects of a greater mutation load could be concealed by parental care by allowing half the lineages to receive post-hatching care, while half did not. We found that inbred lineages from the Full Care populations went extinct more quickly than inbred lineages from the No Care populations-but only when offspring received no post-hatching care. We infer that Full Care lineages carried a greater mutation load, but that the associated deleterious effects on fitness could be overcome if larvae received parental care. We suggest that the increased mutation load caused by parental care increases a population's dependence upon care. This could explain why care is seldom lost once it has evolved.

Dark extinction: the problem of unknown historical extinctions.

The extinction of species before they are discovered and named (dark extinction, DE) is widely inferred as a significant part of species loss in the 'pre-taxonomic' period (approx. 1500-1800 CE) and, to some extent, in the 'taxonomic period' (approx. 1800-present) as well. The discovery of oceanic islands and other pristine habitats by European navigators and the consequent introduction of destructive mammals, such as rats and goats, started a process of anthropogenic extinction. Much ecosystem change happened before systematic scientific recording, so has led to DE. Statistical methods are available to robustly estimate DE in the 'taxonomic period'. For the 'pre-taxonomic period', simple extrapolation can be used. The application of these techniques to world birds, for example, suggests that approximately 56 DEs occurred in the 'taxonomic period' (1800-present) and approximately 180 in the 'pre-taxonomic period' (1500-1800). Targeting collection activities in extinction hotspots, to make sure organisms are represented in collections before their extinction, is one way of reducing the number of extinct species without a physical record (providing that collection efforts do not themselves contribute to species extinction).

Vascular plant extinction in the continental United States and Canada.

Extinction rates are expected to increase during the Anthropocene. Current extinction rates of plants and many animals remain unknown. We quantified extinctions among the vascular flora of the continental United States and Canada since European settlement. We compiled data on apparently extinct species by querying plant conservation databases, searching the literature, and vetting the resulting list with botanical experts. Because taxonomic opinion varies widely, we developed an index of taxonomic uncertainty (ITU). The ITU ranges from A to F, with A indicating unanimous taxonomic recognition and F indicating taxonomic recognition by only a single author. The ITU allowed us to rigorously evaluate extinction rates. Our data suggest that 51 species and 14 infraspecific taxa, representing 33 families and 49 genera of vascular plants, have become extinct in our study area since European settlement. Seven of these taxa exist in cultivation but are extinct in the wild. Most extinctions occurred in the west, but this outcome may reflect the timing of botanical exploration relative to settlement. Sixty-four percent of extinct plants were single-site endemics, and many occurred outside recognized biodiversity hotspots. Given the paucity of plant surveys in many areas, particularly prior to European settlement, the actual extinction rate of vascular plants is undoubtedly much higher than indicated here.

Extinción de las Plantas Vasculares en Canadá y los Estados Unidos Continentales Resumen Se espera que las tasas de extinción aumenten durante el Antropoceno. Todavía desconocemos las tasas de extinción actuales de las plantas y muchos animales. Cuantificamos las tasas de extinción de la flora vascular de los Estados Unidos Continentales y Canadá a partir del asentamiento de los europeos. Recopilamos datos sobre especies aparentemente extintas mediante la consulta de bases de datos sobre conservación, búsquedas en la literatura y el escrutinio de la lista resultante con botánicos expertos. Ya que la opinión taxonómica varía ampliamente, desarrollamos un índice de incertidumbre taxonómica (ITU). La ITU abarca desde la A hasta la F, en donde la A indica un reconocimiento taxonómico unánime y la F indica el reconocimiento taxonómico por un sólo autor. La ITU nos permitió evaluar rigurosamente las tasas de extinción. Nuestros datos sugieren que 51 especies y 14 taxones infraespecíficos, que en conjunto representan a 33 familias y a 49 géneros de plantas vasculares, se han extinguido en nuestra área de estudio desde el asentamiento de los europeos. Siete de estos taxones existen en cultivos, pero se encuentran extintos en vida libre. La mayoría de las extinciones ocurrieron en la parte oeste del área de estudio, aunque este resultado puede reflejar el momento de la exploración botánica en relación con el asentamiento europeo. El 64% de las plantas extintas eran endémicas de un sitio único y muchas tuvieron presencia fuera de los puntos calientes de biodiversidad. Dada la escasez de los censos botánicos en muchas áreas, particularmente previo al asentamiento europeo, la tasa actual de extinción de las plantas vasculares es sin duda mucho más alta de lo que se indica en este estudio.

Extinction risk in extant marine species integrating palaeontological and biodistributional data.

Extinction risk assessments of marine invertebrate species remain scarce, which hinders effective management of marine biodiversity in the face of anthropogenic impacts. To help close this information gap, in this paper we provide a metric of relative extinction risk that combines palaeontological data, in the form of extinction rates calculated from the fossil record, with two known correlates of risk in the modern day: geographical range size and realized thermal niche. We test the performance of this metric-Palaeontological Extinction Risk In Lineages (PERIL)-using survivorship analyses of Pliocene bivalve faunas from California and New Zealand, and then use it to identify present-day hotspots of extinction vulnerability for extant shallow-marine Bivalvia. Areas of the ocean where concentrations of bivalve species with higher PERIL scores overlap with high levels of climatic or anthropogenic stressors should be considered of most immediate concern for both conservation and management.

Application of fundamental equations to species-area theory.

BACKGROUND: Species-area relationship (SAR), endemics-area relationship (EAR) and overlap-area relationship (OAR) are three important concepts in biodiversity study. The application of fundamental equations linking the SAR, EAR and OAR, can enrich the axiomatic framework of the species-area theory and deepen our understanding of the mechanisms of community assembly. RESULTS: Two fundamental equations are derived and extended to power law model and random replacement model of species-area distribution. Several important parameters, including the overlap index and extinction rate, are defined and expressed to enrich the species-area theory. For power law model, both EAR and OAR have three parameters, with one more parameter of the total area than SAR does. The EAR equation is a monotonically increasing function for parameter c and z, and a monotonically decreasing function for parameter A. The extinction rate, with two parameters, is a monotonically increasing function for parameter z, and a monotonically decreasing function for parameter A. The overlap index is a monotonically increasing function for parameter A, and a monotonically decreasing function for parameter z, independent of parameter c. CONCLUSIONS: The general formats of SAR, EAR, OAR, overlap index, overlap rate, sampling rate and extinction rate, are derived and extended to power law model and random replacement model as the axiomatic framework of species-area theory. In addition, if the total area is underestimated, the extinction rate will be overestimated.

Confluence, synnovation, and depauperons in plant diversification.

We review the empirical phylogenetic literature on plant diversification, highlighting challenges in separating the effects of speciation and extinction, in specifying diversification mechanisms, and in making convincing arguments. In recent discussions of context dependence, key opportunities and landscapes, and indirect effects and lag times, we see a distinct shift away from single-point/single-cause 'key innovation' hypotheses toward more nuanced explanations involving multiple interacting causal agents assembled step-wise through a tree. To help crystalize this emerging perspective we introduce the term 'synnovation' (a hybrid of 'synergy' and 'innovation') for an interacting combination of traits with a particular consequence ('key synnovation' in the case of increased diversification rate), and the term 'confluence' for the sequential coming together of a set of traits (innovations and synnovations), environmental changes, and geographic movements along the branches of a phylogenetic tree. We illustrate these concepts using the radiation of Bromeliaceae. We also highlight the generality of these ideas by considering how rate heterogeneity associated with a confluence relates to the existence of particularly species-poor lineages, or 'depauperons.' Many challenges are posed by this re-purposed research framework, including difficulties associated with partial taxon sampling, uncertainty in divergence time estimation, and extinction.

Temperate radiations and dying embers of a tropical past: the diversification of Viburnum.

We used a near-complete phylogeny for the angiosperm clade Viburnum to assess lineage diversification rates, and to examine possible morphological and ecological factors driving radiations. Maximum-likelihood and Bayesian approaches identified shifts in diversification rate and possible links to character evolution. We inferred the ancestral environment for Viburnum and changes in diversification dynamics associated with subsequent biome shifts. Viburnum probably diversified in tropical forests of Southeast Asia in the Eocene, with three subsequent radiations in temperate clades during the Miocene. Four traits (purple fruits, extrafloral nectaries, bud scales and toothed leaves) were statistically associated with higher rates of diversification. However, we argue that these traits are unlikely to be driving diversification directly. Instead, two radiations were associated with the occupation of mountainous regions and a third with repeated shifts between colder and warmer temperate forests. Early-branching depauperate lineages imply that the rare lowland tropical species are 'dying embers' of once more diverse lineages; net diversification rates in Viburnum likely decreased in these tropical environments after the Oligocene. We suggest that 'taxon pulse' dynamics might characterize other temperate plant lineages.

Estimating the normal background rate of species extinction.

A key measure of humanity's global impact is by how much it has increased species extinction rates. Familiar statements are that these are 100-1000 times pre-human or background extinction levels. Estimating recent rates is straightforward, but establishing a background rate for comparison is not. Previous researchers chose an approximate benchmark of 1 extinction per million species per year (E/MSY). We explored disparate lines of evidence that suggest a substantially lower estimate. Fossil data yield direct estimates of extinction rates, but they are temporally coarse, mostly limited to marine hard-bodied taxa, and generally involve genera not species. Based on these data, typical background loss is 0.01 genera per million genera per year. Molecular phylogenies are available for more taxa and ecosystems, but it is debated whether they can be used to estimate separately speciation and extinction rates. We selected data to address known concerns and used them to determine median extinction estimates from statistical distributions of probable values for terrestrial plants and animals. We then created simulations to explore effects of violating model assumptions. Finally, we compiled estimates of diversification-the difference between speciation and extinction rates for different taxa. Median estimates of extinction rates ranged from 0.023 to 0.135 E/MSY. Simulation results suggested over- and under-estimation of extinction from individual phylogenies partially canceled each other out when large sets of phylogenies were analyzed. There was no evidence for recent and widespread pre-human overall declines in diversity. This implies that average extinction rates are less than average diversification rates. Median diversification rates were 0.05-0.2 new species per million species per year. On the basis of these results, we concluded that typical rates of background extinction may be closer to 0.1 E/MSY. Thus, current extinction rates are 1,000 times higher than natural background rates of extinction and future rates are likely to be 10,000 times higher.

Settling down of seasonal migrants promotes bird diversification.

How seasonal migration originated and impacted diversification in birds remains largely unknown. Although migratory behaviour is likely to affect bird diversification, previous studies have not detected any effect. Here, we infer ancestral migratory behaviour and the effect of seasonal migration on speciation and extinction dynamics using a complete bird tree of life. Our analyses infer that sedentary behaviour is ancestral, and that migratory behaviour evolved independently multiple times during the evolutionary history of birds. Speciation of a sedentary species into two sedentary daughter species is more frequent than speciation of a migratory species into two migratory daughter species. However, migratory species often diversify by generating a sedentary daughter species in addition to the ancestral migratory one. This leads to an overall higher migratory speciation rate. Migratory species also experience lower extinction rates. Hence, although migratory species represent a minority (18.5%) of all extant birds, they have a higher net diversification rate than sedentary species. These results suggest that the evolution of seasonal migration in birds has facilitated diversification through the divergence of migratory subpopulations that become sedentary, and illustrate asymmetrical diversification as a mechanism by which diversification rates are decoupled from species richness.

Color polymorphic carnivores have faster speciation rates.

Variation in coat color is a prominent feature in carnivores, thought to be shaped by environmental factors. As new traits could allow populations to occupy novel niches and habitats, color polymorphism may be maintained by balancing selection. Consequently, color polymorphic species may speciate more rapidly and can give rise to monomorphic daughter species. We thus predicted that, within the Carnivora, (i) speciation rate is higher in polymorphic lineages, (ii) divergence between color polymorphic lineages is more recent, and (iii) within closely related groups, polymorphic lineages are ancestral and monomorphic lineages derived. We also tested whether accelerated speciation rates relate to niche breadth, measured by the number of occupied habitats and range size. We collected data of 48 polymorphic and 192 monomorphic carnivore species, and assessed speciation rates using phylogenetic comparative methods. We found that polymorphic carnivores had higher speciation rates (λ1 = 0.29, SD = 0.13) than monomorphic species (λ0 = 0.053, SD = 0.044). Hidden and quantitative state speciation and extinction models inferred that color polymorphism was the main contributing factor, and that niche breadth was not of influence. Therefore, other selective forces than spatial niche segregation, such as predator-prey coevolution, may contribute to color polymorphism in wild carnivores.

Larger brains spur species diversification in birds.

Evidence is accumulating that species traits can spur their evolutionary diversification by influencing niche shifts, range expansions, and extinction risk. Previous work has shown that larger brains (relative to body size) facilitate niche shifts and range expansions by enhancing behavioral plasticity but whether larger brains also promote evolutionary diversification is currently backed by insufficient evidence. We addressed this gap by combining a brain size dataset for >1900 avian species worldwide with estimates of diversification rates based on two conceptually different phylogenetic-based approaches. We found consistent evidence that lineages with larger brains (relative to body size) have diversified faster than lineages with relatively smaller brains. The best supported trait-dependent model suggests that brain size primarily affects diversification rates by increasing speciation rather than decreasing extinction rates. In addition, we found that the effect of relatively brain size on species-level diversification rate is additive to the effect of other intrinsic and extrinsic factors. Altogether, our results highlight the importance of brain size as an important factor in evolution and reinforce the view that intrinsic features of species have the potential to influence the pace of evolution.

Recent Anthropogenic Plant Extinctions Differ in Biodiversity Hotspots and Coldspots.

During the Anthropocene, humans are changing the Earth system in ways that will be detectable for millennia to come [1]. Biologically, these changes include habitat destruction, biotic homogenization, increased species invasions, and accelerated extinctions [2]. Contemporary extinction rates far surpass background rates [3], but they seem remarkably low in plants [4, 5]. However, biodiversity is not evenly distributed, and as a result, extinction rates may vary among regions. Some authors have contentiously argued that novel anthropic habitats and human-induced plant speciation can actually increase regional biodiversity [6, 7]. Here, we report on one of the most comprehensive datasets to date, including regional and global plant extinctions in both biodiversity hotspots (mostly from Mediterranean-type climate regions) and coldspots (mostly from Eurasian countries). Our data come from regions covering 15.3% of the Earth's surface and span over 300 years. With this dataset, we explore the trends, causes, and temporal dynamics of recent plant extinctions. We found more, and faster accrual of, absolute numbers of extinction events in biodiversity hotspots compared to coldspots. Extinction rates were also substantially higher than historical background rates, but recent declines are evident. We found higher levels of taxonomic uniqueness being lost in biodiversity coldspots compared to hotspots. Causes of plant extinctions also showed distinct temporal patterns, with agriculture, invasions, and urbanization being significant drivers in hotspots, while hydrological disturbance was an important driver in coldspots. Overall, plant extinctions over the last three centuries appear to be low, with a recent (post-1990) and steady extinction rate of 1.26 extinctions/year.

Aging asexual lineages and the evolutionary maintenance of sex.

Finite populations of asexual and highly selfing species suffer from a reduced efficacy of selection. Such populations are thought to decline in fitness over time due to accumulating slightly deleterious mutations or failing to adapt to changing conditions. These within-population processes that lead nonrecombining species to extinction may help maintain sex and outcrossing through species level selection. Although inefficient selection is proposed to elevate extinction rates over time, previous models of species selection for sex assumed constant diversification rates. For sex to persist, classic models require that asexual species diversify at rates lower than sexual species; the validity of this requirement is questionable, both conceptually and empirically. We extend past models by allowing asexual lineages to decline in diversification rates as they age, that is nonrecombining lineages "senesce" in diversification rates. At equilibrium, senescing diversification rates maintain sex even when asexual lineages, at young ages, diversify faster than their sexual progenitors. In such cases, the age distribution of asexual lineages contains a peak at intermediate values rather than showing the exponential decline predicted by the classic model. Coexistence requires only that the average rate of diversification in asexuals be lower than that of sexuals.

Extinction as a driver of avian latitudinal diversity gradients.

The role of historical factors in driving latitudinal diversity gradients is poorly understood. Here, we used an updated global phylogeny of terrestrial birds to test the role of three key historical factors-speciation, extinction, and dispersal rates-in generating latitudinal diversity gradients for eight major clades. We fit a model that allows speciation, extinction, and dispersal rates to differ, both with latitude and between the New and Old World. Our results consistently support extinction (all clades had lowest extinction where species richness was highest) as a key driver of species richness gradients across each of eight major clades. In contrast, speciation and dispersal rates showed no consistent latitudinal patterns across replicate bird clades, and thus are unlikely to represent general underlying drivers of latitudinal diversity gradients.