The Natural History Museum repository: Recent submissions
Now showing items 1-20 of 1374
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Boosting biodiversity in school grounds: a theory of changeThe National Education Nature Park aims to involve every nursery, school, and college in England in enhancing the biodiversity on their site, whilst supporting young people’s wellbeing, pro-environmental behaviours, and green skills. Young people gather environmental data using citizen science research, and then through collaboration and collective decision-making, they design and implement their own nature recovery actions. But will this participation in community and citizen science lead to behaviour change and environmental action, and how can we build participants’ sense of agency to take environmental action through our programme? Here, we present our Theory of Change for the Nature Park and the design features of the programme that connect participation in citizen science with achieving two crucial types of change - environmental change in the form of biodiversity gain, and the behaviour change that underpins it.
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Effects of land use and soil properties on taxon richness and abundance of soil assemblagesAbstract: Land‐use change and habitat degradation are among the biggest drivers of aboveground biodiversity worldwide but their effects on soil biodiversity are less well known, despite the importance of soil organisms in developing soil structure, nutrient cycling and water drainage. Combining a global compilation of biodiversity data from soil assemblages collated as part of the PREDICTS project with global data on soil characteristics, we modelled how taxon richness and total abundance of soil organisms have responded to land use. We also estimated the global Biodiversity Intactness Index (BII)—the average abundance and compositional similarity of taxa that remain in an area, compared to a minimally impacted baseline, for soil biodiversity. This is the first time the BII has been calculated for soil biodiversity. Relative to undisturbed vegetation, soil organism total abundance and taxon richness were reduced in all land uses except pasture. Soil properties mediated the response of soil biota, but not in a consistent way across land uses. The global soil BII in cropland is, on average, a third of that originally present. However, in grazed sites the decline is less severe. The BII of secondary vegetation depends on age, with sites with younger growth showing a lower BII than mature vegetation. We conclude that land‐use change has reduced local soil biodiversity worldwide, and this further supports the proposition that soil biota should be considered explicitly when using global models to estimate the state of biodiversity.
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The genome sequence of the false flower beetle, Anaspis frontalis (Linnaeus, 1758)We present a genome assembly from a specimen of <ns5:italic>Anaspis frontalis</ns5:italic> (the false flower beetle; Arthropoda; Insecta; Coleoptera; Scraptiidae). The assembly contains two haplotypes with total lengths of 808.55 megabases and 802.05 megabases. Most of haplotype 1 (95.81%) is scaffolded into 8 chromosomal pseudomolecules, including the X chromosome, while haplotype 2 is a scaffold-level assembly. The mitochondrial genome has also been assembled and is 16.47 kilobases in length.
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The genome sequence of the click beetle, Ampedus sanguinolentus sanguinolentus (Schrank, 1776)We present a genome assembly from a female specimen of <ns3:italic>Ampedus sanguinolentus sanguinolentus</ns3:italic> (click beetle; Arthropoda; Insecta; Coleoptera; Elateridae). The assembly contains two haplotypes with total lengths of 1,574.76 megabases and 1,572.87 megabases. Most of haplotype 1 (97.13%) is scaffolded into 10 chromosomal pseudomolecules, while haplotype 2 is a scaffold-level assembly. The mitochondrial genome has also been assembled and is 15.99 kilobases in length.
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The genome sequence of a longhorn beetle, Rhagium mordax (Degeer, 1775)We present a genome assembly from an individual female specimen of <ns5:italic>Rhagium mordax</ns5:italic> (longhorn beetle; Arthropoda; Insecta; Coleoptera; Cerambycidae). The genome sequence has a total length of 775.60 megabases. Most of the assembly (99.53%) is scaffolded into 10 chromosomal pseudomolecules. The mitochondrial genome has also been assembled and is 16.68 kilobases in length. Gene annotation of this assembly on Ensembl identified 11,937 protein-coding genes.
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Revisiting the genus Bolbosoma Porta, 1908 (Acanthocephala: Polymorphidae): host specificity, phylogeny, and species synonymizationAbstract Background: Acanthocephalans of the genus <jats:italic>Bolbosoma</jats:italic> Porta, 1908 are trophically transmitted parasites that infect marine mammals (mostly cetaceans and less frequently pinnipeds) worldwide. There are 12 species currently considered as valid; however, most records lack information on the maturity stage of the specimens. This, coupled with the scarce phylogenetic information available, hinders a correct understanding of their patterns of host specificity, evolutionary history, and taxonomy. A particularly intriguing case is that of <jats:italic>Bolbosoma vasculosum</jats:italic> (Rudolphi, 1819), which has been frequently reported in odontocetes but rarely as an adult, having been suggested to be synonymous with <jats:italic>Bolbosoma capitatum</jats:italic> (von Linstow, 1880). Methods: We used a comprehensive approach to investigate the concept of <jats:italic>Bolbosoma</jats:italic>. First, we conducted a bibliographic review of records of <jats:italic>Bolbosoma</jats:italic> spp. to clarify which are the final hosts for each species. We paid particular attention to <jats:italic>B. vasculosum</jats:italic>, using morphological and molecular analyses to compare it with <jats:italic>B. capitatum</jats:italic>. Second, we characterized the complete mitochondrial genome of <jats:italic>Bolbosoma balaenae</jats:italic> (Gmelin, 1790), <jats:italic>Bolbosoma turbinella</jats:italic> (Diesing, 1851), <jats:italic>B. capitatum</jats:italic>, and <jats:italic>B. vasculosum</jats:italic>. Then, we reconstructed the phylogenetic relationships of <jats:italic>Bolbosoma</jats:italic> spp. and related taxa using full mitochondrial genomes (or only <jats:italic>cox1</jats:italic> when full mitogenomes were unavailable) and nuclear ribosomal genes (18S and 28S). Results: Bolbosoma</jats:italic> spp. exhibit high specificity for cetaceans, with no confirmed records of adult specimens in other host groups. Within this genus, <jats:italic>B. vasculosum</jats:italic> appears to be conspecific with <jats:italic>B. capitatum</jats:italic> based on both morphological and molecular evidence. This species shows high affinity to odontocetes, while the remaining species are specific to mysticetes. Phylogenetic analyses showed strong support for the monophyly of <jats:italic>Bolbosoma</jats:italic> spp., which appeared as sister taxa to <jats:italic>Corynosoma</jats:italic> spp. and <jats:italic>Andracantha</jats:italic> spp.. The resulting topology aligns with the patterns of specificity indicated by host records, revealing two distinct clades for species specific to odontocetes and mysticetes, respectively. Conclusions: The phylogenetic relationships obtained support the hypothesis that the association of <jats:italic>Bolbosoma</jats:italic> spp. with cetaceans originated through a host-switching event from aquatic birds.
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The genome sequence of the false flower beetle, Anaspis regimbarti Schilsky, 1895We present a genome assembly from a specimen of <ns3:italic>Anaspis regimbarti</ns3:italic> (the false flower beetle; Arthropoda; Insecta; Coleoptera; Scraptiidae). The genome sequence has a total length of 457.61 megabases. Most of the assembly (99.89%) is scaffolded into 8 chromosomal pseudomolecules, including the X sex chromosome. The mitochondrial genome has also been assembled and is 16.39 kilobases in length.
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Redescription of Mesoschendyla javanica (Attems, 1907) and its first records from Borneo (Chilopoda, Geophilomorpha, Schendylidae)The geophilomorph Mesoschendyla javanica (Attems, 1907) was originally described from a small number of males collected from bat guano in Tjompea (Ciampea), Java. Subsequently, no additional material was identified. The type series remained the only specimens belonging to this genus known from Asia. Mesoschendyla javanica is re-discovered 118 years after its original description amongst centipedes collected from soil cores taken during the 1977–1978 Royal Geographical Society Gunung Mulu Expedition to Sarawak (Borneo, East Malaysia) and is deposited in the collection of the Natural History Museum (London). The new material comprising 49 specimens, amongst which are the first known females, is described and illustrated, shedding light on intraspecific morphological variation. The syntypes and sole previously available specimens are redescribed and illustrated, completing the summary original description of the species. Ecological and biogeographical notes are provided for Sarawak specimens.
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Phytochemical, Antioxidant and Anticancer Properties of Ayusip (Vaccinium uliginosum L.) from Benguet Province, PhilippinesTraditional medicinal knowledge remains an important source of healthcare for many communities in the Philippines, particularly in rural and Indigenous areas where access to modern treatments is limited. In this context, Vaccinium uliginosum (locally known as Ayusip), a plant used by indigenous peoples in Tublay, Benguet Province, has drawn interest due to its reported therapeutic applications for ailments such as cough, flu, and even cancer. This study investigates the cytotoxic properties of Ayusip, aiming to provide scientific validation for its traditional use and assess its potential as a source of bioactive compounds for cancer treatment. Leaves of V. uliginosum were collected, identified, and subjected to methanol extraction, with further partitioning using n-hexane and ethyl acetate. Phytochemical analysis indicated high levels of flavonoids, saponins, steroids, and tannins. Quantification showed significant phenolic (24.58±8.11 GAE mg/g) and flavonoid (121.38±14.04 QE mg/g) content. Antioxidant testing (DPPH assay) revealed strong activity (97.86±1.33%) at 25 mg/mL. The cytotoxicity of V. uliginosum was assessed against normal embryonic lung fibroblasts (MRC-5) and cancer cell lines HCT-116 (colorectal) and A549 (lung adenocarcinoma) using the MTT assay. Results showed that only the n-hexane extract was toxic to MRC-5 cells at high concentrations, while methanol and n-hexane extracts were highly cytotoxic to HCT-116 cells. In A549 cells, only n-hexane extract exhibited cytotoxicity. These findings suggest that V. uliginosum has promising cytotoxic effects, particularly at higher concentrations, with notable activity against HCT-116 cells. Interestingly, its cytotoxicity is biphasic: high concentrations induce maximum toxicity, whereas lower concentrations may encourage cell proliferation. Overall, this study offers insights into the cytotoxic potential of V. uliginosum, a traditionally used plant among Benguet’s IPs, highlighting its potential medicinal value.
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First record of Calyptotheca alexandriensis (Cheilostomatida, Lanceoporidae) from Ras Juddi (Pasni) Makran coast, Northern Arabian SeaThe available information regarding bryozoans in coastal areas of Pakistan is limited. This research paper presents the first record of Calyptotheca alexandriensis from Ras Juddi (Pasni) along the Makran coast, and the second record globally of this species. This species was first reported in the Eastern Harbor of Alexandria, Egypt, as a distinctive deep orange erect foliaceous bryozoan and was observed abundantly on various hard substrates, such as rocks, ropes, metal pipes supporting marina piers, and ship hulls. The discovery adds a new record of the family, Lanceoporidae, and genus, Calyptotheca, to the bryozoan fauna of Pakistan, expanding its known distribution to the Northern Arabian Sea. The present specimens were collected during December 2021–October 2022 and subjected to detailed taxonomic analysis using light microscopy and scanning electron microscopy (SEM). This study contributes to the understanding of bryozoan biodiversity in the region and highlights the potential for further discoveries along the unexplored coastal areas of Pakistan.
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Phylogenetic origin of an insect fauna at the boundary of the Palaearctic and Oriental realms: Evidence from ‘site‐based’ mitogenomicsAbstract Aim: Knowledge of taxonomy and species distributions in highly diverse lineages of invertebrates remains too incomplete for the study of biogeographical patterns at the global scale. This limits the understanding of processes leading to the formation and maintenance of boundaries between major biogeographical realms. Site‐based metagenomic approaches may provide an alternative source of data for inference of historical processes of in situ speciation and interchange among biogeographical regions. We applied the methodology to explore the historical biogeography of the Qinling Mountains at the boundary of the Oriental and Palaearctic regions. Location: The focal area in the Qinling Mountains; sites in Borneo, Panama, and Spain, representing the Oriental, Neotropical and Palaearctic faunas. Time Period: Contemporary. Major Taxa Studied: Chrysomelidae, a group of ~40,000 species with worldwide distribution. Methods: Mass‐trapped insects were subjected to deep Illumina DNA sequencing and mitochondrial genome assembly, followed by phylogenetic analysis and historical biogeographical inference on a dated tree. The methodology produced 622 mitogenomes (304 newly sequenced) each representing a morphologically distinct species. Results: Ancestral area reconstruction revealed that the Qinling fauna is a composite of distinct clades that, respectively, have Oriental or Palaearctic ancient origins, while in situ speciation and local clade formation were limited. The global diversity of Chrysomelidae consists of regionalised deep clades at the level of the major zoogeographic realms, which remain recognisable where these realms abut in the Qinling mountains. Yet, the Qinling fauna is clearly distinct from the source areas although local lineage ages are generally <20 Ma. Main Conclusions: The Qinling fauna is a composite of clades that have either Oriental or Palaearctic origin, indicating the important role of immigration at the realm boundary. Global site‐based phylogenetic analyses based on metagenomic sequencing can address previously intractable questions about the historical biogeography of poorly studied groups even without a fully developed taxonomy.
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Community hydrodynamics created ecological opportunity in Ediacaran shallow marine ecosystemsAbstract: The “second wave” of Ediacaran evolution (∼558–548 Ma) was characterized by the appearance of macroscopic organisms in shallow marine settings, where they formed communities with high morphological and ecological diversity, including new and more complex modes of life. Based on analogy with modern marine ecosystems, these early shallow water communities could have substantially modified local hydrodynamic conditions and influenced resource availability, but we know very little about how they interacted with their fluid environment at larger spatial scales. Here, we use computational fluid dynamics to investigate the hydrodynamics of different shallow marine Ediacaran communities based on fossil surfaces from Russia and South Australia. Our results reveal considerable hydrodynamic variability among these communities, ranging from unobstructed flow, to enhanced mixing, to very low in-canopy flow. This variability represents a noticeable shift from the more conserved hydrodynamic conditions reconstructed for older Ediacaran communities from deep water settings. The variation in how shallow marine Ediacaran communities affected local hydrodynamics could have given rise to notable differences in the distribution of crucial water-borne resources such as organic carbon and oxygen. We therefore hypothesize that increasing variability in community hydrodynamics was an important source of habitat heterogeneity during the late Ediacaran. On long timescales, this heterogeneity may have helped sculpt ecological opportunity, fostering the radiation of animals.
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Molecular phylogenies provide insights into the evolutionary relationships of the Spirurida (Nematoda), with special emphasis on the superfamily PhysalopteroideaAbstract Background Nematodes of the order Spirurida are of significant veterinary, medical, and economic importance. However, current knowledge of the phylogenetic relationships within the order is far from comprehensive. Moreover, the monophyly of the Physalopteroidea/Physalopteridae, and the phylogenetic relationships of its three component subfamilies, remain uncertain due to inadequate sequence data. Methods The nuclear small ribosomal subunit (18S rRNA) and large ribosomal subunit (28S rRNA), plus the complete mitochondrial genomes of two physalopterid species, Thubunaea pudica (Thubunaeinae) and Abbreviata varani (Physalopterinae), are presented for the first time. Phylogenetic analyses of the Spirurida were performed using maximum likelihood and Bayesian inference on the basis of different concatenated datasets involving the most comprehensive subfamily-level taxon sampling of the superfamily Physalopteroidea to date to provide an initial understanding of the evolutionary relationships of major superfamilies within the order, with special emphasis on the Physalopteroidea/Physalopteridae. Results The complete mitogenomes of T. pudica and A. varani are 13,645 bp and 13,730 bp in length, which both contain 36 genes and belong to the GA9 type gene arrangement. Molecular phylogenies based on different datasets all support a close affinity between the superfamilies Camallanoidea and Dracunculoidea. Our phylogenetic results also showed that the representatives of the Physalopteridae did not form a monophyletic group. The representative of the subfamily Proleptinae clustered together with the tetramerid species Crassicauda magna, and the representative of the subfamily Thubunaeinae formed a sister relationship with species of the subfamily Physalopterinae. Conclusions The mitogenome of T. pudica is the first for the subfamily Thubunaeinae, and that of A. varani is also the first for the genus Abbreviata . Molecular phylogenetic results suggest that the subfamily Proleptinae should be elevated to full family level, i.e., Proleptidae stat. nov., and that the genus Crassicauda (Habronematoidea: Tetrameridae) should be assigned to Proleptidae stat. nov. Our results also indicate that the Physalopteridae sensu stricto currently comprises two subfamilies, the Thubunaeinae and the Physalopterinae, and that the genus Skrjabinoptera should be transferred from the Physalopterinae to the Thubunaeinae.
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On the natural history of West African Rock Dove Columba gymnocyclaNamed in 1856, West African Rock Dove Columba livia gymnocycla differs markedly in appearance from other Rock Dove subspecies. Recent DNA analyses have revealed that gymnocycla is also genetically very different from Columba livia, warranting species status for this taxon. The species was never abundant in its small distribution, and only a small number of specimens are present in museum collections. A search for the species in Senegambia in 2024 was unsuccessful and C. gymnocycla may have already vanished from much or all of its distribution.
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Taxonomic identities of Psittacula cyanopygia Souancé, 1856, Pachycephala clio Wallace, 1863, and Lagonosticta brunneiceps Sharpe, 1890, fixed through designation of lectotypes held in the Natural History Museum, TringDuring ongoing work in the Natural History Museum, Tring, bird collections to identify and segregate syntype material, three cases have been identified where it is necessary to objectively define the relevant taxa. The first of these is the Mexican Parrotlet Forpus cyanopygius (Souancé, 1856), of which the male and female syntypes are identifiable as different species, with the female being a specimen of Cobalt-rumped Parrotlet F. xanthopterygius. Although some authors have suggested that the type material of F. cyanopygius is from the Tres Marías Islands, off western Mexico, and thus referrable to F. cyanopygius insularis, we demonstrate that the male syntype (now lectotype) can be confidently identified as being from the Mexican mainland, obviating the need for any nomenclatural revision. The second case involves Pachycephala clio Wallace, 1863, currently treated as a subspecies of Moluccan Whistler P. macrorhyncha, which was described from specimens collected in the Sula Islands and on Buru. Subsequently, Pachycephala melanura buruensis E. Hartert, 1899, now also a subspecies of P. macrorhyncha, was named from specimens collected exclusively on Buru, but without unambiguously restricting Wallace's nomen to the Sula Islands. The final case pertains to Lagonosticta brunneiceps Sharpe, 1890, which is currently treated as a subspecies of Red-billed Firefinch L. senegala. Specimens of Sharpe's type series originated from a large part of the distribution of L. senegala and are representative of multiple currently accepted subspecies. Therefore, lectotype designations are offered to fix the identity on the universally understood taxonomic concept associated with all of these taxa.
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Cortical and trabecular bone structure of the hominoid capitateAbstract: Morphological variation in the hominoid capitate has been linked to differences in habitual locomotor activity due to its importance in movement and load transfer at the midcarpal joint proximally and carpometacarpal joints distally. Although the shape of bones and their articulations are linked to joint mobility, the internal structure of bones has been shown experimentally to reflect, at least in part, the loading direction and magnitude experienced by the bone. To date, it is uncertain whether locomotor differences among hominoids are reflected in the bone microarchitecture of the capitate. Here, we apply a whole‐bone methodology to quantify the cortical and trabecular architecture (separately and combined) of the capitate across bipedal (modern<jats:italic>Homo sapiens</jats:italic>), knuckle‐walking (<jats:italic>Pan paniscus</jats:italic>,<jats:italic>Pan troglodytes</jats:italic>,<jats:italic>Gorilla</jats:italic>sp.), and suspensory (<jats:italic>Pongo</jats:italic>sp.) hominoids (<jats:italic>n</jats:italic> = 69). It is hypothesized that variation in bone microarchitecture will differentiate these locomotor groups, reflecting differences in habitual postures and presumed loading force and direction. Additionally, it is hypothesized that trabecular and cortical architecture in the proximal and distal regions, as a result of being part of mechanically divergent joints proximally and distally, will differ across these portions of the capitate. Results indicate that the capitate of knuckle‐walking and suspensory hominoids is differentiated from bipedal<jats:italic>Homo</jats:italic>primarily by significantly thicker distal cortical bone. Knuckle‐walking taxa are further differentiated from suspensory and bipedal taxa by more isotropic trabeculae in the proximal capitate. An allometric analysis indicates that size is not a significant determinate of bone variation across hominoids, although sexual dimorphism may influence some parameters within<jats:italic>Gorilla</jats:italic>. Results suggest that internal trabecular and cortical bone is subjected to different forces and functional adaptation responses across the capitate (and possibly other short bones). Additionally, while separating trabecular and cortical bone is normal protocol of current whole‐bone methodologies, this study shows that when applied to carpals, removing or studying the cortical bone separately potentially obfuscates functionally relevant signals in bone structure.</jats:p>
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Eocene Loranthaceae pollen pushes back divergence ages for major splits in the familyBackground: We revisit the palaeopalynological record of Loranthaceae, using pollen ornamentation to discriminate lineages and to test molecular dating estimates for the diversification of major lineages. Methods: Fossil Loranthaceae pollen from the Eocene and Oligocene are analysed and documented using scanning-electron microscopy. These fossils were associated with molecular-defined clades and used as minimum age constraints for Bayesian node dating using different topological scenarios. Results: The fossil Loranthaceae pollen document the presence of at least one extant root-parasitic lineage (Nuytsieae) and two currently aerial parasitic lineages (Psittacanthinae and Loranthinae) by the end of the Eocene in the Northern Hemisphere. Phases of increased lineage diversification (late Eocene, middle Miocene) coincide with global warm phases. Discussion: With the generation of molecular data becoming easier and less expensive every day, neontological research should re-focus on conserved morphologies that can be traced through the fossil record. The pollen, representing the male gametophytic generation of plants and often a taxonomic indicator, can be such a tracer. Analogously, palaeontological research should put more effort into diagnosing Cenozoic fossils with the aim of including them into modern systematic frameworks.
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Multi-rate Poisson tree processes for single-locus species delimitation under maximum likelihood and Markov chain Monte CarloAbstract Motivation: In recent years, molecular species delimitation has become a routine approach for quantifying and classifying biodiversity. Barcoding methods are of particular importance in large-scale surveys as they promote fast species discovery and biodiversity estimates. Among those, distance-based methods are the most common choice as they scale well with large datasets; however, they are sensitive to similarity threshold parameters and they ignore evolutionary relationships. The recently introduced “Poisson Tree Processes” (PTP) method is a phylogeny-aware approach that does not rely on such thresholds. Yet, two weaknesses of PTP impact its accuracy and practicality when applied to large datasets; it does not account for divergent intraspecific variation and is slow for a large number of sequences. Results: We introduce the multi-rate PTP (mPTP), an improved method that alleviates the theoretical and technical shortcomings of PTP. It incorporates different levels of intraspecific genetic diversity deriving from differences in either the evolutionary history or sampling of each species. Results on empirical data suggest that mPTP is superior to PTP and popular distance-based methods as it, consistently yields more accurate delimitations with respect to the taxonomy (i.e., identifies more taxonomic species, infers species numbers closer to the taxonomy). Moreover, mPTP does not require any similarity threshold as input. The novel dynamic programming algorithm attains a speedup of at least five orders of magnitude compared to PTP, allowing it to delimit species in large (meta-) barcoding data. In addition, Markov Chain Monte Carlo sampling provides a comprehensive evaluation of the inferred delimitation in just a few seconds for millions of steps, independently of tree size. Availability and Implementation: mPTP is implemented in C and is available for download at http://github.com/Pas-Kapli/mptp under the GNU Affero 3 license. A web-service is available at http://mptp.h-its.org. Supplementary information: Supplementary data are available at Bioinformatics online.
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Species relationships and divergence times in beeches: new insights from the inclusion of 53 young and old fossils in a birth–death clock modelThe fossilized birth–death (FBD) model can make use of information contained in multiple fossils representing the same clade, and we here apply this model to infer divergence times in beeches (genus <jats:italic>Fagus</jats:italic>), using 53 fossils and nuclear sequences for all nine species. We also apply FBD dating to the fern clade Osmundaceae, with about 12 living species and 36 fossils. <jats:italic>Fagus</jats:italic> nuclear sequences cannot be aligned with those of other Fagaceae, and we therefore use Bayes factors to choose among alternative root positions. The crown group of <jats:italic>Fagus</jats:italic> is dated to 53 (62–43) Ma; divergence of the sole American species to 44 (51–39) Ma and divergence between Central European <jats:italic>F. sylvatica</jats:italic> and Eastern Mediterranean <jats:italic>F. orientalis</jats:italic> to 8.7 (20–1.8) Ma, unexpectedly old. The FBD model can accommodate fossils as sampled ancestors or as extinct or unobserved lineages; however, this makes its raw output, which shows all fossils on short or long branches, problematic to interpret. We use hand-drawn depictions and a bipartition network to illustrate the uncertain placements of fossils. Inferred speciation and extinction rates imply approximately 5× higher evolutionary turnover in <jats:italic>Fagus</jats:italic> than in Osmundaceae, fitting a hypothesized low turnover in plants adapted to low-nutrient conditions. This article is part of the themed issue ‘Dating species divergences using rocks and clocks’.













