The origin of eukaryotes and rise in complexity were synchronous with the rise in oxygen.

The origin of eukaryotes was among the most important events in the history of life, spawning a new evolutionary lineage that led to all complex multicellular organisms. However, the timing of this event, crucial for understanding its environmental context, has been difficult to establish. The fossil and biomarker records are sparse and molecular clocks have thus far not reached a consensus, with dates spanning 2.1-0.91 billion years ago (Ga) for critical nodes. Notably, molecular time estimates for the last common ancestor of eukaryotes are typically hundreds of millions of years younger than the Great Oxidation Event (GOE, 2.43-2.22 Ga), leading researchers to question the presumptive link between eukaryotes and oxygen. We obtained a new time estimate for the origin of eukaryotes using genetic data of both archaeal and bacterial origin, the latter rarely used in past studies. We also avoided potential calibration biases that may have affected earlier studies. We obtained a conservative interval of 2.2-1.5 Ga, with an even narrower core interval of 2.0-1.8 Ga, for the origin of eukaryotes, a period closely aligned with the rise in oxygen. We further reconstructed the history of biological complexity across the tree of life using three universal measures: cell types, genes, and genome size. We found that the rise in complexity was temporally consistent with and followed a pattern similar to the rise in oxygen. This suggests a causal relationship stemming from the increased energy needs of complex life fulfilled by oxygen.

The colonisation of Madagascar by land-bound vertebrates.

Despite discussions extending back almost 160 years, the means by which Madagascar's iconic land vertebrates arrived on the island remains the focus of active debate. Three options have been considered: vicariance, range expansion across land bridges, and dispersal over water. The first assumes that a group (clade/lineage) occupied the island when it was connected with the other Gondwana landmasses in the Mesozoic. Causeways to Africa do not exist today, but have been proposed by some researchers for various times in the Cenozoic. Over-water dispersal could be from rafting on floating vegetation (flotsam) or by swimming/drifting. A recent appraisal of the geological data supported the idea of vicariance, but found nothing to justify the notion of past causeways. Here we review the biological evidence for the mechanisms that explain the origins of 28 of Madagascar's land vertebrate clades [two other lineages (the geckos Geckolepis and Paragehyra) could not be included in the analysis due to phylogenetic uncertainties]. The podocnemid turtles and typhlopoid snakes are conspicuous for they appear to have arisen through a deep-time vicariance event. The two options for the remaining 26 (16 reptile, five land-bound-mammal, and five amphibian), which arrived between the latest Cretaceous and the present, are dispersal across land bridges or over water. As these would produce very different temporal influx patterns, we assembled and analysed published arrival times for each of the groups. For all, a 'colonisation interval' was generated that was bracketed by its 'stem-old' and 'crown-young' tree-node ages; in two instances, the ranges were refined using palaeontological data. The synthesis of these intervals for all clades, which we term a colonisation profile, has a distinctive shape that can be compared, statistically, to various models, including those that assume the arrivals were focused in time. The analysis leads us to reject the various land bridge models (which would show temporal concentrations) and instead supports the idea of dispersal over water (temporally random). Therefore, the biological evidence is now in agreement with the geological evidence, as well as the filtered taxonomic composition of the fauna, in supporting over-water dispersal as the mechanism that explains all but two of Madagascar's land-vertebrate groups.

Discovering research articles containing evolutionary timetrees by machine learning.

MOTIVATION: Timetrees depict evolutionary relationships between species and the geological times of their divergence. Hundreds of research articles containing timetrees are published in scientific journals every year. The TimeTree (TT) project has been manually locating, curating and synthesizing timetrees from these articles for almost two decades into a TimeTree of Life, delivered through a unique, user-friendly web interface (timetree.org). The manual process of finding articles containing timetrees is becoming increasingly expensive and time-consuming. So, we have explored the effectiveness of text-mining approaches and developed optimizations to find research articles containing timetrees automatically. RESULTS: We have developed an optimized machine learning system to determine if a research article contains an evolutionary timetree appropriate for inclusion in the TT resource. We found that BERT classification fine-tuned on whole-text articles achieved an F1 score of 0.67, which we increased to 0.88 by text-mining article excerpts surrounding the mentioning of figures. The new method is implemented in the TimeTreeFinder (TTF) tool, which automatically processes millions of articles to discover timetree-containing articles. We estimate that the TTF tool would produce twice as many timetree-containing articles as those discovered manually, whose inclusion in the TT database would potentially double the knowledge accessible to a wider community. Manual inspection showed that the precision on out-of-distribution recently published articles is 87%. This automation will speed up the collection and curation of timetrees with much lower human and time costs. AVAILABILITY AND IMPLEMENTATION: https://github.com/marija-stanojevic/time-tree-classification. SUPPLEMENTARY INFORMATION: Supplementary data are available at Bioinformatics online.

Completing a molecular timetree of apes and monkeys.

The primate infraorder Simiiformes, comprising Old and New World monkeys and apes, includes the most well-studied species on earth. Their most comprehensive molecular timetree, assembled from thousands of published studies, is found in the TimeTree database and contains 268 simiiform species. It is, however, missing 38 out of 306 named species in the NCBI taxonomy for which at least one molecular sequence exists in the NCBI GenBank. We developed a three-pronged approach to expanding the timetree of Simiiformes to contain 306 species. First, molecular divergence times were searched and found for 21 missing species in timetrees published across 15 studies. Second, untimed molecular phylogenies were searched and scaled to time using relaxed clocks to add four more species. Third, we reconstructed ten new timetrees from genetic data in GenBank, allowing us to incorporate 13 more species. Finally, we assembled the most comprehensive molecular timetree of Simiiformes containing all 306 species for which any molecular data exists. We compared the species divergence times with those previously imputed using statistical approaches in the absence of molecular data. The latter data-less imputed times were not significantly correlated with those derived from the molecular data. Also, using phylogenies containing imputed times produced different trends of evolutionary distinctiveness and speciation rates over time than those produced using the molecular timetree. These results demonstrate that more complete clade-specific timetrees can be produced by analyzing existing information, which we hope will encourage future efforts to fill in the missing taxa in the global timetree of life.

TimeTree 5: An Expanded Resource for Species Divergence Times.

We present the fifth edition of the TimeTree of Life resource (TToL5), a product of the timetree of life project that aims to synthesize published molecular timetrees and make evolutionary knowledge easily accessible to all. Using the TToL5 web portal, users can retrieve published studies and divergence times between species, the timeline of a species' evolution beginning with the origin of life, and the timetree for a given evolutionary group at the desired taxonomic rank. TToL5 contains divergence time information on 137,306 species, 41% more than the previous edition. The TToL5 web interface is now ADA-compliant and mobile-friendly, a result of comprehensive source code refactoring. TToL5 also offers programmatic access to species divergence times and timelines through an application programming interface, which is accessible at timetree.temple.edu/api. TToL5 is publicly available at timetree.org.

A global reptile assessment highlights shared conservation needs of tetrapods.

Comprehensive assessments of species' extinction risks have documented the extinction crisis1 and underpinned strategies for reducing those risks2. Global assessments reveal that, among tetrapods, 40.7% of amphibians, 25.4% of mammals and 13.6% of birds are threatened with extinction3. Because global assessments have been lacking, reptiles have been omitted from conservation-prioritization analyses that encompass other tetrapods4-7. Reptiles are unusually diverse in arid regions, suggesting that they may have different conservation needs6. Here we provide a comprehensive extinction-risk assessment of reptiles and show that at least 1,829 out of 10,196 species (21.1%) are threatened-confirming a previous extrapolation8 and representing 15.6 billion years of phylogenetic diversity. Reptiles are threatened by the same major factors that threaten other tetrapods-agriculture, logging, urban development and invasive species-although the threat posed by climate change remains uncertain. Reptiles inhabiting forests, where these threats are strongest, are more threatened than those in arid habitats, contrary to our prediction. Birds, mammals and amphibians are unexpectedly good surrogates for the conservation of reptiles, although threatened reptiles with the smallest ranges tend to be isolated from other threatened tetrapods. Although some reptiles-including most species of crocodiles and turtles-require urgent, targeted action to prevent extinctions, efforts to protect other tetrapods, such as habitat preservation and control of trade and invasive species, will probably also benefit many reptiles.

Limitations of Phylogenomic Data Can Drive Inferred Speciation Rate Shifts.

Biodiversity analyses of phylogenomic timetrees have produced many high-profile examples of shifts in the rate of speciation across the tree of life. Temporally correlated events in ecology, climate, and biogeography are frequently invoked to explain these rate shifts. In a re-examination of 15 genomic timetrees and 25 major published studies of the pattern of speciation through time, we observed an unexpected correlation between the timing of reported rate shifts and the information content of sequence alignments. Here, we show that the paucity of sequence variation and insufficient species sampling in phylogenomic data sets are the likely drivers of many inferred speciation rate shifts, rather than the proposed biological explanations. Therefore, data limitations can produce predictable but spurious signals of rate shifts even when speciation rates may be similar across taxa and time. Our results suggest that the reliable detection of speciation rate shifts requires the acquisition and assembly of long phylogenomic alignments with near-complete species sampling and accurate estimates of species richness for the clades of study.

A new species of frog from the Caribbean island of Montserrat (Eleutherodactylidae, Eleutherodactylus).

I describe a new species of frog, Eleutherodactylus montserratae sp. nov., from Montserrat, previously confused with E. johnstonei Barbour. It is native to that island and has been introduced, and established, on other Lesser Antillean islands west of 62 degrees west longitude including Anguilla, St. Martin/St. Maarten, St. Barthélemy, Saba, St. Eustatius, St. Christopher, and Nevis, as well as Bermuda. Previous chromosome and molecular studies have shown that populations from these islands are distinct from E. johnstonei, a species occurring east of 62 degrees west longitude in the Lesser Antilles, and elsewhere, but could not identify diagnostic morphological differences. Here, I show that the new species differs morphologically in being smaller (males and females) and having a proportionately larger tympanum, a wider head, and greater separation between the nostrils. Both species have been introduced within and outside of the Caribbean region and will likely expand their ranges in the future.

A new genus and species of Caribbean forest lizard (Diploglossidae; Celestinae) from southern Hispaniola.

A new genus and species of Caribbean forest lizard of the subfamily Celestinae Schools & Hedges are described. This new taxon is known only from two small adjacent keys in the Laguna de Oviedo of the Parque Nacional Jaragua in the Barahona Peninsula, Dominican Republic, at the southernmost region of the Caribbean island of Hispaniola. The genus Guarocuyus gen. nov. is genetically more closely related to the clade containing Celestus Gray, Comptus Schools & Hedges, and Panolopus Cope; nevertheless, we compare it to all celestine genera. Morphologically, it differs from other celestines by having an interdigital web on three toes and by having the widest ear opening. Additionally, the species Guarocuyus jaraguanus sp. nov. has some ecological attributes that when combined, appear to be unique, including nocturnal habits, a semiprehensile tail, and a facultatively arboreal lifestyle. We note sexual dimorphism in the new species and in two other celestines, Caribicus darlingtoni (Cochran) and the poorly known Celestus macrotus Thomas & Hedges, and report a range extension of the latter species into the Dominican Republic. We also discuss several scale topography conditions considered of taxonomic value for the group.

A new semifossorial snake of the genus Arrhyton (Squamata: Dipsadidae) from eastern Cuba, with taxonomic comments on other species.

The genus Arrhyton is endemic to Cuba, with eight currently known species. A new species, Arrhyton albicollum sp. nov., is described from the karst areas of Gibara, Holgun Province. The new species is closely related to A. redimitum, another eastern species, according to morphology and molecular phylogeny. It differs from all other species of the genus by having a conspicuous, white nuchal band, a contrasting black pattern on the head, a gray-colored body with faint stripes, and a high number of ventral and subcaudal scales. Additional comments on other species include new locality records and information on morphology and relationships.

Phylogenetics, classification, and biogeography of the Neotropical forest lizards (Squamata, Diploglossidae).

Lizards of the family Diploglossidae occur in moist, tropical forests of Middle America, South America, and Caribbean islands. Our analyses based on new molecular and morphological data indicate that the widely distributed genera Celestus Gray, 1839 and Diploglossus Wiegmann, 1834 are paraphyletic. We restrict the former to Caribbean islands and the latter to South America and Caribbean islands. We assign species in Middle America, formerly placed in Celestus and Diploglossus, to Advenus gen. nov., Mesoamericus gen. nov., and Siderolamprus Cope, 1861. We assign species on Caribbean islands, formerly placed in Celestus, to Caribicus gen. nov., Comptus gen. nov., Celestus, Panolopus Cope, 1862, Sauresia Gray, 1852, and Wetmorena Cochran, 1927. Our phylogenetic tree supports three major clades in the family: Celestinae subfam. nov. (Advenus gen. nov., Caribicus gen. nov., Comptus gen. nov., Celestus, Panolopus, Sauresia, and Wetmorena), Diploglossinae (Diploglossus and Ophiodes Wagler, 1828), and Siderolamprinae subfam. nov. (Mesoamericus gen. nov. and Siderolamprus). Our timetree indicates that the diploglossid lineage originated in the early Cenozoic and established three major centers of diversification in the Americas: Middle America (siderolamprines and one celestine), South America (diploglossines), and Caribbean islands (celestines and diploglossines). The majority of threatened species are on Caribbean islands, with the major threats being deforestation and predation by the introduced mongoose. Molecular and morphological data indicate that there are many undescribed species in this family of lizards.

Large-Scale Phylogenomic Analyses Reveal the Monophyly of Bryophytes and Neoproterozoic Origin of Land Plants.

The relationships among the four major embryophyte lineages (mosses, liverworts, hornworts, vascular plants) and the timing of the origin of land plants are enigmatic problems in plant evolution. Here, we resolve the monophyly of bryophytes by improving taxon sampling of hornworts and eliminating the effect of synonymous substitutions. We then estimate the divergence time of crown embryophytes based on three fossil calibration strategies, and reveal that maximum calibration constraints have a major effect on estimating the time of origin of land plants. Moreover, comparison of priors and posteriors provides a guide for evaluating the optimal calibration strategy. By considering the reliability of fossil calibrations and the influences of molecular data, we estimate that land plants originated in the Precambrian (980-682 Ma), much older than widely recognized. Our study highlights the important contribution of molecular data when faced with contentious fossil evidence, and that fossil calibrations used in estimating the timescale of plant evolution require critical scrutiny.

A morphological and molecular revision of lizards of the genus Marisora Hedges amp; Conn (Squamata: Mabuyidae) from Central America and Mexico, with descriptions of four new species.

The skink genus Marisora ranges from Mexico to northern South America and occurs on some islands in the Caribbean Sea. We conducted a revision of the genus Marisora from Mexico and Central America, using new morphological and molecular data, and find support for the five previously described species (Marisora alliacea, M.aurulae, M. brachypoda, M. magnacornae, and M. roatanae) and describe four new species: Marisora lineola sp. nov., M. aquilonaria sp. nov., M. syntoma sp. nov., and M. urtica sp. nov. We show that two species previously known only from Central American islands, M. magnacornae and M. roatanae, also occur on the adjacent mainland and that two species recently placed in Alinea belong to this evolutionary clade: Marisora berengerae n. comb. and Marisora pergravis n. comb. Together with M. falconensis and M. unimarginata, these 13 species of Marisora arose mostly in the Pliocene and are largely allopatric but are sympatric and nearly sympatric at several locations in Central America where they maintain their morphological and genetic distinctiveness.

Species diversity as a surrogate for conservation of phylogenetic and functional diversity in terrestrial vertebrates across the Americas.

Preserving the evolutionary history and ecological functions that different species embody, in addition to species themselves, is a growing concern for conservation. Recent studies warn that conservation priority regions identified using species diversity differ from those based on phylogenetic or functional diversity. However, spatial mismatches in conservation priority regions need not indicate low surrogacy among these dimensions in conservation planning. Here, we use data for 10,213 terrestrial vertebrate species across the Americas to evaluate surrogacy; that is, the proportion of phylogenetic or functional diversity represented in conservation plans targeting species. We find that most conservation plans targeting species diversity also represent phylogenetic and functional diversity well, despite spatial mismatches in the priority regions identified by each plan. However, not all phylogenetic and functional diversity is represented within species-based plans, with the highest-surrogacy conservation strategy depending on the proportion of land area included in plans. Our results indicate that targeting species diversity could be sufficient to preserve much of the phylogenetic and functional dimensions of biodiversity in terrestrial vertebrates of the Americas. Incorporating phylogenetic and functional data in broad-scale conservation planning may not always be necessary, especially when the cost of doing so is high.

Haiti's biodiversity threatened by nearly complete loss of primary forest.

Tropical forests hold most of Earth's biodiversity. Their continued loss through deforestation and agriculture is the main threat to species globally, more than disease, invasive species, and climate change. However, not all tropical forests have the same ability to sustain biodiversity. Those that have been disturbed by humans, including forests previously cleared and regrown (secondary growth), have lower levels of species richness compared with undisturbed (primary) forests. The difference is even greater considering extinctions that will later emanate from the disturbance (extinction debt). Here, we find that Haiti has less than 1% of its original primary forest and is therefore among the most deforested countries. Primary forest has declined over three decades inside national parks, and 42 of the 50 highest and largest mountains have lost all primary forest. Our surveys of vertebrate diversity (especially amphibians and reptiles) on mountaintops indicates that endemic species have been lost along with the loss of forest. At the current rate, Haiti will lose essentially all of its primary forest during the next two decades and is already undergoing a mass extinction of its biodiversity because of deforestation. These findings point to the need, in general, for better reporting of forest cover data of relevance to biodiversity, instead of "total forest" as defined by the United Nation's Food and Agricultural Organization. Expanded detection and monitoring of primary forest globally will improve the efficiency of conservation measures, inside and outside of protected areas.

Molecular evidence for the paraphyly of Scolecophidia and its evolutionary implications.

The phylogenetic relationships between the three main clades of worm snakes remain controversial. This question is, however, crucial to elucidate the origin of the successful snake radiation, as these burrowing and miniaturized wormlike organisms represent the earliest branching clades within the snake tree. The present molecular phylogenetic study, intended to minimize the amount of missing data, provides fully resolved inter-subfamilial relationships among Typhlopidae. It also brings robust evidence that worm snakes (Scolecophidia) are paraphyletic, with the scolecophidian family Anomalepididae recovered with strong support as sister clade to the 'typical snakes' (Alethinophidia). Ancestral state reconstructions applied to three different traits strongly associated to a burrowing life-style (scolecoidy, absence of retinal cones and microstomy) provide results in favour of a burrowing origin of snakes, and suggest that worm snakes might be the only extant fossorial representatives of the primordial snake incursion towards an underground environment.