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Premise of research. Fern fronds are common in Late Paleozoic and Mesozoic strata. Large tri- or multipinnate fronds are mainly preserved as impression/compressions, while anatomically preserved sp...

One of the two lineages that constitute the crown group Testudines, Pan-Pleurodira, experienced an important radiation during the Early Cretaceous, from the Barremian or earlier. The origin of the two lineages that form the crown Pleurodira (i.e. Pan-Chelidae and Pelomedusoides) is related to this radiation. Pelomedusoides diversified in northern Gondwana in a relatively short period of geological time. The African Aptian fossiliferous region of Gadoufaoua (central Niger) is one of the most interesting fossiliferous areas for the study of the early diversity and evolution of Pelomedusoides, this being where the oldest African identification of the synchronic and sympatric presence of several pleurodiran taxa is recognised. At least two lineages of Pelomedusoides are represented in Gadoufaoua, one being the extinct Araripemydidae. The second lineage had been recognised as related to Pelomedusidae or Podocnemidoidea, both clades representing part of the current biodiversity. A form from Gadoufaoua, hitherto poorly known and preliminarily presented almost 40 years ago, and currently determined as aff. Platycheloides sp., belongs to this second lineage. Its detailed study is performed here. It is defined as Francemys gadoufaouaensis gen. et sp. nov., constituting one of the few nominated pan-pleurodiran taxa (i.e., about a dozen species) currently recognised in the Lower Cretaceous record. It is identified as being closely related to Podocnemidoidea, the clade that groups the common ancestor and all of the descendants of the abundant and diverse extinct bothremydids and extant podocnemidids.

Astragalus, a highly diverse genus of flowering plants with its highest center of diversity in West Asia, is a classic example of rapid species-level radiation and adaptation to a diversity of habitats throughout the world. We examined the historical biogeography of Astragalus using molecular dating and ancestral area reconstruction to understand how past climate changes, geographical patterns and transition in life history have provoked diversification of Astragalus. Our results suggest that Astragalus probably originated during the middle Miocene in West Asia, underwent rapid diversification, subsequently and repeatedly expanded its range in the Mediterranean region, and later to North America through West Europe. This distribution range was also extended toward central and eastern Asia from the Middle Miocene to Pleistocene. Several climatic and geological processes during the Miocene-Pliocene may be implicated in the diversification of the major Astragalus clades. In particular, the annual lineages, which are important elements in the Mediterranean flora of Africa and Europe and in the deserts of southwest to central Asia, have arisen in response to progressing aridity from the late Miocene onwards (between 8.6 Ma and 2.98 Ma). Diversification rate analyses indicate three rapid and recent diversification events, one at c. 11 Ma in the clade that groups most of the Astragalus s.s. (all except the Ophiocarpus sister lineage), one at c. 5 Ma in the crown group of the Hypoglottis clade, including herbaceous annual and perennial species, and the most recent one at c. 3 Ma in the spiny cushion forming Astracantha clade. Our study highlights the complexity of processes and factors shaping diversifications in Astragalus; a complex interaction among climatic modifications providing opportunities for diversification and likely coincident with the evolution of key morphological and physiological adaptations.

Background and Aims Whole-genome duplication (WGD) events are considered important driving forces of diversification. At least 11 out of 52 Brassicaceae tribes had independent mesopolyploid WGDs followed by diploidization processes. However, the association between mesopolyploidy and subsequent diversification is equivocal. Herein we show the results from a family-wide diversification analysis on Brassicaceae, and elaborate on the hypothesis that polyploidization per se is a fundamental driver in Brassicaceae evolution. Methods We established a time-calibrated chronogram based on whole plastid genomes comprising representative Brassicaceae taxa and published data spanning the entire Rosidae clade. This allowed us to set multiple calibration points and anchored various Brassicaceae taxa for subsequent downstream analyses. All major splits among Brassicaceae lineages were used in BEAST analyses of 48 individually analysed tribes comprising 2101 taxa in total using the internal transcribed spacers of nuclear ribosomal DNA. Diversification patterns were investigated on these tribe-wide chronograms using BAMM and were compared with family-wide data on genome size variation and species richness. Key Results Brassicaceae diverged 29.9 million years ago (Mya) during the Oligocene, and the majority of tribes started diversification in the Miocene with an average crown group age of about 12.5 Mya. This matches the cooling phase right after the Mid Miocene climatic optimum. Significant rate shifts were detected in 12 out of 52 tribes during the Mio- and Pliocene, decoupled from preceding mesopolyploid WGDs. Among the various factors analysed, the combined effect of tribal crown group age and net diversification rate (speciation minus extinction) is likely to explain sufficiently species richness across Brassicaceae tribes. Conclusions The onset of the evolutionary splits among tribes took place under cooler and drier conditions. Pleistocene glacial cycles may have contributed to the maintenance of high diversification rates. Rate shifts are not consistently associated with mesopolyploid WGD. We propose, therefore, that WGDs in general serve as a constant ‘pump’ for continuous and high species diversification.

We describe a nearly complete, three‐dimensionally preserved skull of a new albatross species from the late Pliocene (3.0–3.4 million years ago) Tangahoe Formation of New Zealand. Aldiomedes angustirostris, n. gen. et sp. has only about 90% of the length of the skull of the smallest extant albatross and is the geologically youngest record of a small‐sized albatross known to date. The new species is characterized by a mediolaterally compressed beak, which is not found in any living albatross. The small size and some cranial features of A. angustirostris indicate that, in spite of its comparatively young geological age, the new species was not part of crown group Diomedeidae. We hypothesize that A. angustirostris was more piscivorous than extant albatrosses, which predominantly feed on squid. The reasons for the extinction of smaller‐sized albatrosses are elusive but may be related to changes in seabird fauna during the Pliocene epoch, which witnessed the radiation of various non‐procellariiform seabird groups.

In crown group tetrapods, individual digits are homologized in relation to a pentadactyl ground plan. However, testing hypotheses of digit homology is challenging because it is unclear whether digits represent distinct and conserved gene regulatory states. Here we show dramatic evolutionary dynamism in the gene expression profiles of digits, challenging the notion that five digits have conserved developmental identities across amniotes. Transcriptomics shows diversity in the patterns of gene expression differentiation of digits, although the anterior-most digit of the pentadactyl limb has a unique, conserved expression profile. Further, we identify a core set of transcription factors that are differentially expressed among the digits of amniote limbs; their spatial expression domains, however, vary between species. In light of these results, we reevaluate the frame shift hypothesis of avian wing evolution and conclude only the identity of the anterior-most digit has shifted position, suggesting a 1,3,4 digit identity in the bird wing., The homology of digits across amniotes is debated. Here, the authors compare the developmental transcriptomes of digits across five divergent amniotes and show high evolutionary dynamism in expression profiles, with conservation of a distinct developmental identity only in the anterior-most digit.

The crown wasps, family Stephanidae, are generally believed to occupy a distinguished position as putative relicts of the earliest-diverging lineage of apocritan Hymenoptera (e.g., Sharkey et al., 2012; Mao et al., 2015). More recent analyses have cast some confusion over this hypothesis, with the family instead appearing closer to the Evanioidea or even Trigonalyoidea (Peters et al., 2017; Tang et al., 2019). From most analyses it is clear that the family extends well into the Cretaceous, with crown-group Stephanidae estimated to have appeared by at least the Early Cretaceous and a purported ghost-stem lineage extending into the Early Jurassic or even latest Triassic (Tang et al., 2019). At least parts of such a hypothesis are consistent with the number of mid-Cretaceous fossils representing a variety of crown wasps, including species of both the plesiomorphic subfamily Schlettereriinae as well as putative Stephaninae (Engel & Grimaldi, 2004; Engel et al., 2013; Engel & Huang, 2017; Li et al., 2017). Unfortunately, while such fossil occurrences are of considerable interest, the total available record of fossil crown wasps is poor, with most species documented from the Palaeogene (Engel, 2005; Engel & Ortega-Blanco, 2008), and hitherto only four species from the Late Cretaceous. Given the potentially long gap between the first divergence of the lineage and the appearance of the crown group (Tang et al., 2019), it is precisely for such a group that early diverging stem groups would be of considerable value in resolving relationships and documenting the appearance of apomorphies within the clade. Extensive study of Early Cretaceous and Jurassic deposits for stem-group Stephanidae is necessary in order to provide direct evidence into the early evolution of this critical family of the Euhymenoptera.

Micropeltidaceae species are flyspeck fungi which have been subjected to few systematic studies. We re-examined 27 genera which were accepted in the Micropeltidaceae and re-described them based on herbaria materials and protologues. Based on morphology and phylogenetic investigations, we transfer Micropeltidaceae to a new order, Micropeltidales (Lecanoromycetes). Genera with bluish or greenish upper walls (Dictyopeltella, Dictyothyriella, Dictyothyrina, Dictyothyrium, Haplopeltheca, Micropeltis, Scolecopeltidium and Stomiopeltopsis) are accepted in the new taxonomic concept for Micropeltidaceae. A molecular clock approach estimated the divergence time of the Micropeltidaceae crown group at 130 (165–104) Mya, which also supports its rank as an order (diverging from 220–100 Mya). The evolutionary histories between Micropeltidaceae species and host plants are interpreted by cophylogenetic analyses calibrated by their divergence times. The result indicates that the diversification of Angiospermae (130–80 Mya) fosters the formation of genera of Micropeltidaceae mainly via cospeciation events, and this codivergent period would be an important reference when establishing generic boundaries of epifoliar fungi.

Fungi are crucial components of modern ecosystems. They may have had an important role in the colonization of land by eukaryotes, and in the appearance and success of land plants and metazoans1–3. Nevertheless, fossils that can unambiguously be identified as fungi are absent from the fossil record until the middle of the Palaeozoic era4,5. Here we show, using morphological, ultrastructural and spectroscopic analyses, that multicellular organic-walled microfossils preserved in shale of the Grassy Bay Formation (Shaler Supergroup, Arctic Canada), which dates to approximately 1,010–890 million years ago, have a fungal affinity. These microfossils are more than half a billion years older than previously reported unambiguous occurrences of fungi, a date which is consistent with data from molecular clocks for the emergence of this clade6,7. In extending the fossil record of the fungi, this finding also pushes back the minimum date for the appearance of eukaryotic crown group Opisthokonta, which comprises metazoans, fungi and their protist relatives8,9. Morphological, ultrastructural and spectroscopic analyses identify a fungal affinity for microfossils in shale from Arctic Canada, which pushes back the date for this kingdom to 1,010–890 million years ago.

Floristic similarities between European and Asian mountain ranges have long been recognized, and the hypothesis that European mountain plant taxa immigrated from Asian mountain areas has been confirmed by several molecular phylogenetic analyses. Callianthemum contains ca. 14 species, of which ca. 11 are distributed in Asia and three in Europe. A molecular phylogenetic analysis of the genus using ITS and four plastid DNA sequences (matK, rpL32–trnL intergenic spacer, trnL–trnF intergenic spacer, trnV–ndhC intergenic spacer) revealed that the genus reached Europe twice, with C. coriandrifolium representing one, and C. anemonoides and C. kernerianum a second lineage. Support for C. anemonoides and C. kernerianum as sister species is weak. The crown group ages of the C. coriandrifolium lineage (median 2.2 million years) and of the C. anemonoides/C. kernerianum lineage (median 1.62 million years) are similar and place their immigration to Europe in the Quaternary. Analysis of climatic data shows that C. coriandrifolium grows in colder climate than C. anemonoides/C. kernerianum and that the climatic niches of the three European species are different considering all climatic variables analyzed.

Significance Jawless, boneless, and virtually without fossil record, hagfish have long escaped systematists’ grip on their place among other fish. Yet their systematic resolution is critical to define vertebrates as a clade. Here we report an unequivocal fossil hagfish from the Cretaceous Mediterranean. Using this fossil to calibrate the evolutionary history of the group, our analysis supports hagfish and lampreys as sister groups, which likely diverged from one another in early Paleozoic times. As a result, vertebrates have a deep dichotomy, where some fossil jawless vertebrates sit closer to hagfish and lampreys than to jawed vertebrates. We showed that morphology-based analysis converged onto molecular inferences when characters are coded nonindependently, providing a case study for morphological–molecular conflicts in animal phylogeny., Hagfish depart so much from other fishes anatomically that they were sometimes considered not fully vertebrate. They may represent: (i) an anatomically primitive outgroup of vertebrates (the morphology-based craniate hypothesis); or (ii) an anatomically degenerate vertebrate lineage sister to lampreys (the molecular-based cyclostome hypothesis). This systematic conundrum has become a prominent case of conflict between morphology- and molecular-based phylogenies. To date, the fossil record has offered few insights to this long-branch problem or the evolutionary history of hagfish in general, because unequivocal fossil members of the group are unknown. Here, we report an unequivocal fossil hagfish from the early Late Cretaceous of Lebanon. The soft tissue anatomy includes key attributes of living hagfish: cartilages of barbels, postcranial position of branchial apparatus, and chemical traces of slime glands. This indicates that the suite of characters unique to living hagfish appeared well before Cretaceous times. This new hagfish prompted a reevaluation of morphological characters for interrelationships among jawless vertebrates. By addressing nonindependence of characters, our phylogenetic analyses recovered hagfish and lampreys in a clade of cyclostomes (congruent with the cyclostome hypothesis) using only morphological data. This new phylogeny places the fossil taxon within the hagfish crown group, and resolved other putative fossil cyclostomes to the stem of either hagfish or lamprey crown groups. These results potentially resolve the morphological–molecular conflict at the base of the Vertebrata. Thus, assessment of character nonindependence may help reconcile morphological and molecular inferences for other major discords in animal phylogeny.

The consequences of the Cretaceous–Paleogene (K–Pg) boundary (KPB) mass extinction for the evolution of plant diversity remain poorly understood, even though evolutionary turnover of plant lineages at the KPB is central to understanding assembly of the Cenozoic biota. The apparent concentration of whole genome duplication (WGD) events around the KPB may have played a role in survival and subsequent diversification of plant lineages. To gain new insights into the origins of Cenozoic biodiversity, we examine the origin and early evolution of the globally diverse legume family (Leguminosae or Fabaceae). Legumes are ecologically (co-)dominant across many vegetation types, and the fossil record suggests that they rose to such prominence after the KPB in parallel with several well-studied animal clades including Placentalia and Neoaves. Furthermore, multiple WGD events are hypothesized to have occurred early in legume evolution. Using a recently inferred phylogenomic framework, we investigate the placement of WGDs during early legume evolution using gene tree reconciliation methods, gene count data and phylogenetic supernetwork reconstruction. Using 20 fossil calibrations we estimate a revised timeline of legume evolution based on 36 nuclear genes selected as informative and evolving in an approximately clock-like fashion. To establish the timing of WGDs we also date duplication nodes in gene trees. Results suggest either a pan-legume WGD event on the stem lineage of the family, or an allopolyploid event involving (some of) the earliest lineages within the crown group, with additional nested WGDs subtending subfamilies Papilionoideae and Detarioideae. Gene tree reconciliation methods that do not account for allopolyploidy may be misleading in inferring an earlier WGD event at the time of divergence of the two parental lineages of the polyploid, suggesting that the allopolyploid scenario is more likely. We show that the crown age of the legumes dates to the Maastrichtian or early Paleocene and that, apart from the Detarioideae WGD, paleopolyploidy occurred close to the KPB. We conclude that the early evolution of the legumes followed a complex history, in which multiple auto- and/or allopolyploidy events coincided with rapid diversification and in association with the mass extinction event at the KPB, ultimately underpinning the evolutionary success of the Leguminosae in the Cenozoic. [Allopolyploidy; Cretaceous–Paleogene (K–Pg) boundary; Fabaceae, Leguminosae; paleopolyploidy; phylogenomics; whole genome duplication events], info:eu-repo/semantics/published

Understanding the biogeographical and diversification processes explaining current diversity patterns of subcosmopolitan-distributed groups is challenging. We aimed at disentangling the historical biogeography of the subcosmopolitan liverwort genus Lejeunea with estimation of ancestral areas of origin and testing if sexual system and palaeotemperature variations can be factors of diversification. We assembled a dense taxon sampling for 120 species sampled throughout the geographical distribution of the genus. Lejeunea diverged from its sister group after the Paleocene-Eocene boundary (52.2 Ma, 95% credibility intervals 50.1–54.2 Ma), and the initial diversification of the crown group occurred in the early to middle Eocene (44.5 Ma, 95% credibility intervals 38.5–50.8 Ma). The DEC model indicated that (1) Lejeunea likely originated in an area composed of the Neotropics and the Nearctic, (2) dispersals through terrestrial land bridges in the late Oligocene and Miocene allowed Lejeunea to colonize the Old World, (3) the Boreotropical forest covering the northern regions until the late Eocene did not facilitate Lejeunea dispersals, and (4) a single long-distance dispersal event was inferred between the Neotropics and Africa. Biogeographical and diversification analyses show the Miocene was an important period when Lejeunea diversified globally. We found slight support for higher diversification rates of species with both male and female reproductive organs on the same individual (monoicy), and a moderate positive influence of palaeotemperatures on diversification. Our study shows that an ancient origin associated with a dispersal history facilitated by terrestrial land bridges and not long-distance dispersals are likely to explain the subcosmopolitan distribution of Lejeunea. By enhancing the diversification rates, monoicy likely favoured the colonisations of new areas, especially in the Miocene that was a key epoch shaping the worldwide distribution.

The eusporangiate marattialean ferns represent an ancient radiation with a rich fossil record but limited modern diversity in the tropics. The long evolutionary history without close extant relatives has confounded studies of the phylogenetic origin, rooting and timing of marattialean ferns. Here we present new complete plastid genomes of six marattialean species and compiled a plastid genome dataset representing all of the currently accepted marattialean genera. We further supplemented this dataset by compiling a large dataset of mitochondrial genes and a phenotypic data matrix covering both extant and extinct representatives of the lineage. Our phylogenomic and total-evidence analyses corroborated the postulated position of marattialean ferns as the sister to leptosporangiate ferns, and the position of Danaea as the sister to the remaining extant marattialean genera. However, our results provide new evidence that Christensenia is sister to Marattia and that M. cicutifolia actually belongs to Eupodium. The apparently highly reduced rate of molecular evolution in marattialean ferns provides a challenge for dating the key phylogenetic events with molecular clock approaches. We instead applied a parsimony-based total-evidence dating approach, which suggested a Triassic age for the extant crown group. The modern distribution can best be explained as mainly resulting from vicariance following the breakup of Pangaea and Gondwana. We resolved the fossil genera Marattiopsis, Danaeopsis and Qasimia as members of the monophyletic family Marattiaceae, and the Carboniferous genera Sydneia and Radstockia as the monophyletic sister of all other marattialean ferns.

Background Ecdysozoa are the moulting protostomes, including arthropods, tardigrades, and nematodes. Both the molecular and fossil records indicate that Ecdysozoa is an ancient group originating in the terminal Proterozoic, and exceptional fossil biotas show their dominance and diversity at the beginning of the Phanerozoic. However, the nature of the ecdysozoan common ancestor has been difficult to ascertain due to the extreme morphological diversity of extant Ecdysozoa, and the lack of early diverging taxa in ancient fossil biotas. Results Here we re-describe Acosmia maotiania from the early Cambrian Chengjiang Biota of Yunnan Province, China and assign it to stem group Ecdysozoa. Acosmia features a two-part body, with an anterior proboscis bearing a terminal mouth and muscular pharynx, and a posterior annulated trunk with a through gut. Morphological phylogenetic analyses of the protostomes using parsimony, maximum likelihood and Bayesian inference, with coding informed by published experimental decay studies, each placed Acosmia as sister taxon to Cycloneuralia + Panarthropoda—i.e. stem group Ecdysozoa. Ancestral state probabilities were calculated for key ecdysozoan nodes, in order to test characters inferred from fossils to be ancestral for Ecdysozoa. Results support an ancestor of crown group ecdysozoans sharing an annulated vermiform body with a terminal mouth like Acosmia, but also possessing the pharyngeal armature and circumoral structures characteristic of Cambrian cycloneuralians and lobopodians. Conclusions Acosmia is the first taxon placed in the ecdysozoan stem group and provides a constraint to test hypotheses on the early evolution of Ecdysozoa. Our study suggests acquisition of pharyngeal armature, and therefore a change in feeding strategy (e.g. predation), may have characterised the origin and radiation of crown group ecdysozoans from Acosmia-like ancestors.

The segmented trapdoor spiders (Liphistiidae) are the sole surviving family of the suborder Mesothelae, which forms the sister lineage to all other living spiders. Liphistiids have retained a number of plesiomorphic traits and their present-day distribution is limited to East and Southeast Asia. Studying this group has the potential to shed light on the deep evolutionary history of spiders, but the phylogeny and divergence times of the family have not been resolved with confidence. We performed phylogenomic and molecular dating analyses of 2765 ultraconserved element loci from 185 liphistiid taxa. Our analyses show that the crown group of Liphistiidae appeared in the mid-Cretaceous at 102 Ma (95% credibility interval 92–113 Ma), but it was not until the Neogene that much of the diversification within the family occurred in mainland Southeast and East Asia. This diversification was coincident with tectonic events such as the extension of the East Asian continental margin, as well as geological upheavals in Indochina induced by the collision between India and Asia. Our study highlights the important role of major tectonic events in shaping the evolutionary history, present-day diversity, and geographical distribution of mesothele and liphistiid spiders. [biogeography; concatenation; Liphistiidae; molecular dating; summary coalescent; UCEs.]

Despite significant advances in invertebrate phylogenomics over the past decade, the higher-level phylogeny of Pycnogonida (sea spiders) remains elusive. Due to the inaccessibility of some small-bodied lineages, few phylogenetic studies have sampled all sea spider families. Previous efforts based on a handful of genes have yielded unstable tree topologies. Here, we inferred the relationships of 89 sea spider species using targeted capture of the mitochondrial genome, 56 conserved exons, 101 ultraconserved elements, and 3 nuclear ribosomal genes. We inferred molecular divergence times by integrating morphological data for fossil species to calibrate 15 nodes in the arthropod tree of life. This integration of data classes resolved the basal topology of sea spiders with high support. The enigmatic family Austrodecidae was resolved as the sister group to the remaining Pycnogonida and the small-bodied family Rhynchothoracidae as the sister group of the robust-bodied family Pycnogonidae. Molecular divergence time estimation recovered a basal divergence of crown group sea spiders in the Ordovician. Comparison of diversification dynamics with other marine invertebrate taxa that originated in the Paleozoic suggests that sea spiders and some crustacean groups exhibit resilience to mass extinction episodes, relative to mollusk and echinoderm lineages. Deutsche Forschungsgemeinschaft BR5039/3-1 National Science Foundation ANT-0551969, ANT-0440577 Ministerio de Economía y Competitividad POL2006-06399/CGL, CTM2012-39350- C02-01 Antarctic Science AA3010 National Science Foundation IOS-1552610

Although dissimilar in their overall appearance and life habits, the praying mantises (Mantodea) and cockroaches (Blattodea, including their eusocial relatives, the termites [Isoptera]) are grouped within the clade Dictyoptera, based on – among other significant characteristics – the laying of eggs in a compound structure called an ootheca. The origin of the Dictyoptera and the currently recognized taxa within is, however, a controversial topic among entomologists. This has resulted from disparities in the divergence age estimates obtained from phylogenetic analyses based on molecular data together with the limited and controversial fossil evidence attributable to these groups. Here, we report two new oothecae ichnospecies found in a Carnian (237 to 227 mya. lowermost Upper Triassic) deposit from Argentina. Morphological comparisons and Scanning Electron Microscope and X-ray Energy Dispersive Spectroscopy analyses of fossil and extant oothecae of mantises and cockroaches were performed in an attempt to solve their systematic placement within Dictyoptera and fossil allies, such as †Alienoptera. In addition to being the earliest known record of oothecae, this discovery moves the origin of this specialized reproductive strategy back by 100 million years. As direct fossil evidence, these specimens provide an important calibration and reference point that can inform future research on the origins and timing of diversification of the Dictyoptera. Fil: Cariglino, Barbara. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Lara, Maria Belén. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste. Centro de Ecología Aplicada del Litoral. Universidad Nacional del Nordeste. Centro de Ecología Aplicada del Litoral; Argentina Fil: Zavattieri, Ana Maria. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mendoza. Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales. Provincia de Mendoza. Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales. Universidad Nacional de Cuyo. Instituto Argentino de Nivología, Glaciología y Ciencias Ambientales; Argentina

International audience; The anatomy of sharks, rays, and chimaeras (chondrichthyans) is crucial to understanding the evolution of the cranial system in vertebrates due to their position as the sister group to bony fishes (osteichthyans). Strikingly different arrangements of the head in the two constituent chondrichthyan groups—holocephalans and elasmobranchs—have played a pivotal role in the formation of evolutionary hypotheses targeting major cranial structures such as the jaws and pharynx. However, despite the advent of digital dissections as a means of easily visualizing and sharing the results of anatomical studies in three dimensions, information on the musculoskeletal systems of the chondrichthyan head remains largely limited to traditional accounts, many of which are at least a century old. Here, we use synchrotron tomographic data to carry out a digital dissection of a holocephalan and an elasmobranch widely used as model species: the elephantfish, Callorhinchus milii, and the small‐spotted catshark, Scyliorhinus canicula. We describe and figure the skeletal anatomy of the head, labial, mandibular, hyoid, and branchial cartilages in both taxa as well as the muscles of the head and pharynx. In Callorhinchus, we make several new observations regarding the branchial musculature, revealing several previously unreported or ambiguously characterized muscles, likely homologous to their counterparts in the elasmobranch pharynx. We also identify a previously unreported structure linking the pharyngohyal of Callorhinchus to the neurocranium. Finally, we review what is known about the evolution of chondrichthyan cranial muscles from their fossil record and discuss the implications for muscle homology and evolution, broadly concluding that the holocephalan pharynx is likely derived from a more elasmobranch‐like form which is plesiomorphic for the chondrichthyan crown group. This dataset has great potential as a resource, particularly for researchers using these model species for zoological research, functional morphologists requiring models of musculature and skeletons, as well as for palaeontologists seeking comparative models for extinct taxa.

Chondrichthyes (Elasmobranchii and Holocephali) are distinguished by their largely cartilaginous endoskeleton that comprises an uncalcified core overlain by a mineralised layer; in the Elasmobranchii (sharks, skates, rays) this mineralisation takes the form of calcified polygonal tiles known as tesserae. In recent years, these skeletal tissues have been described in ever increasing detail in sharks and rays but those of Holocephali (chimaeroids) have been less well-described, with conflicting accounts as to whether or not tesserae are present. During embryonic ontogeny in holocephalans, cervical vertebrae fuse to form a structure called the synarcual. The synarcual mineralises early and progressively, anteroposteriorly and dorsoventrally, and therefore presents a good skeletal structure in which to observe mineralised tissues in this group. Here we describe the development and mineralisation of the synarcual in an adult and stage 36 elephant shark embryo (Callorhinchus milii). Small, discrete, but irregular blocks of cortical mineralisation are present in stage 36, similar to what has been described recently in embryos of other chimaeroid taxa such asHydrolagus, while inCallorhinchusadults, the blocks of mineralisation have become more irregular, but remain small. This differs from fossil members of the holocephalan crown group (Edaphodon), as well as from stem group holocephalans (e.g., Symmorida,Helodus, Iniopterygiformes), where tessellated cartilage is present, with tesserae being notably larger than inCallorhinchusand showing similarities to elasmobranch tesserae, for example with respect to polygonal shape.

The evolutionary history of Mesozoic mammaliaformes is well studied. Although the backbone of their phylogeny is well resolved, the placement of ecologically specialized groups has remained uncertain. Functional and developmental covariation has long been identified as an important source of phylogenetic error, yet combining incongruent morphological characters altogether is currently a common practice when reconstructing phylogenetic relationships. Ignoring incongruence may inflate the confidence in reconstructing relationships, particularly for the placement of highly derived and ecologically specialized taxa, such as among australosphenidans (particularly, crown monotremes), haramiyidans, and multituberculates. The alternative placement of these highly derived clades can alter the taxonomic constituency and temporal origin of the mammalian crown group. Based on prior hypotheses and correlated homoplasy analyses, we identified cheek teeth and shoulder girdle character complexes as having a high potential to introduce phylogenetic error. We showed that incongruence among mandibulodental, cranial, and postcranial anatomical partitions for the placement of the australosphenidans, haramiyids, and multituberculates could largely be explained by apparently non-phylogenetic covariance from cheek teeth and shoulder girdle characters. Excluding these character complexes brought agreement between anatomical regions and improved the confidence in tree topology. These results emphasize the importance of considering and ameliorating major sources of bias in morphological data, and we anticipate that these will be valuable for confidently integrating morphological and molecular data in phylogenetic and dating analyses.

Ladybirds (family Coccinellidae) are one of the most diverse groups of beetles and globally comprise over 6000 species. Despite their scientific and economic significance, the taxonomy of Coccinellidae remains unstable, and we still know little about their evolutionary history. By using a small number of genes, previous phylogenetic analyses have not reliably resolved the relationships among major ladybird lineages. In this study, we sequenced 94 nuclear protein-coding genes for 214 species of Coccinellidae and 14 outgroups, covering 90 genera and 35 tribes. We found that nucleotide compositional heterogeneity is present among ladybird tribes so that phylogenetic inference at the amino acid level is more reliable than at the DNA level. Based on the maximum likelihood analyses of the amino acid dataset, we recognize three subfamilies in Coccinellidae: Microweiseinae, Monocoryninae stat. nov., and Coccinellinae. The subfamily relationships are strongly supported as (Microweiseinae, (Monocoryninae stat. nov., Coccinellinae)). The tribes of ladybirds are mostly monophyletic, except Ortaliini, Sticholotidini, Scymnini, and Coccidulini. The phylogenetic relationships among tribes of Coccinellinae are still not well resolved, with many nodes weakly supported. Our divergence time analysis suggests that the crown group of extant lady beetles arose in the Early Cretaceous ~ 143 million years ago (Mya) and experienced a rapid diversification during the Late Cretaceous (120–70 Mya). We hypothesize that the boom of angiosperms in the Late Cretaceous promoted the diversification of herbivorous Sternorrhyncha insects, especially aphids, which in turn drove the rapid radiation of predatory lady beetles. In summary, our work provides a comprehensive time-calibrated phylogeny of Coccinellidae that provides a sound framework for revising their classification and understanding the origin of their biodiversity.

From a functional standpoint, the humerus is a key element in the skeleton of vertebrates as it is the forelimb&rsquo, s bone that connects with the pectoral girdle. In most birds, the humerus receives both the forces exerted by the main flight muscles and the aerodynamical stresses exerted upon the wing during locomotion. Despite this functional preeminence, broad scale studies of the morphological disparity of the humerus in the crown group of birds (Neornithes) are lacking. Here, we explore the variation in shape of the humeral outline in modern birds and its evolutionary relationship with size and the evolution of different functional regimes, including several flight strategies, wing propelled diving and complete loss of wing locomotory function. Our findings suggest that most neornithines evolved repeatedly towards a general humeral morphology linked with functional advantages related with more efficient flapping. Lineages evolving high-stress locomotion such as hyperaeriality (e.g., swifts), hovering (e.g., hummingbirds) and wing-propelled diving (e.g., penguins) greatly deviate from this general trend, each exploring different morphologies. Secondarily flightless birds deviate to a lesser degree from their parent clades in humeral morphology likely as a result of the release from constraints related with wing-based locomotion. Furthermore, these taxa show a different allometric trend that flighted birds. Our results reveal that the constraints of aerial and aquatic locomotion are main factors shaping the macroevolution of humeral morphology in modern birds.

Allioideae (e.g., chives, garlics, onions) comprises three mainly temperate tribes: Allieae (800 species from the northern hemisphere), Gilliesieae (80 South American species), and Tulbaghieae (26 Southern African species). We reconstructed the phylogeny of Allioideae (190 species plus 257 species from Agapanthoideae and Amaryllidoideae) based on ITS, matK, ndhF, and rbcL to investigate its historical biogeography and karyotype evolution using newly generated cytomolecular data for Chilean Gilliesieae genera Gethyum, Miersia, Solaria, and Speea. The crown group of Allioideae diversified ∼62 Mya supporting a Gondwanic origin for the subfamily and vicariance as the cause of the intercontinental disjunction of the tribes. Our results support the hypothesis of the Indian tectonic plate carrying Allieae to northern hemisphere (‘out-of-India’ hypothesis). The colonization of the northern hemisphere (∼30 Mya) is correlated with a higher diversification rate in Allium associated to stable x = 8, increase of polyploidy and the geographic expansion in Europe and North America. Tulbaghieae presented x = 6, but with numerical stability (2n = 12). In contrast, the tribe Gilliesieae (x = 6) varied considerably in genome size (associated with Robertsonian translocations), rDNA sites distribution and chromosome number. Our data indicate that evolutionary history of Allioideae tribes is linked to distinct trends of karyotype evolution.

Several unique properties of echinoids (sea urchins) make them useful for exploring macroevolutionary dynamics, including their remarkable fossil record that can be incorporated into explicit phylogenetic hypotheses. However, this potential cannot be exploited without a robust resolution of the echinoid tree of life. We revisit the phylogeny of crown group Echinoidea using both the largest phylogenomic dataset compiled for the clade, as well as a large-scale morphological matrix with a dense fossil sampling. We also gather a new compendium of both tip and node age constraints, allowing us to combine phylogenomic, morphological and stratigraphic data using a total-evidence dating approach. For this, we develop a novel method for subsampling phylogenomic datasets that selects loci with high phylogenetic signal, low systematic biases and enhanced clock-like behavior. Our approach restructure much of the higher-level phylogeny of echinoids, and demonstrates that combining different data sources increases topological accuracy. We are able to resolve multiple alleged conflicts between molecular and morphological datasets, such as the position of Echinothurioida and Echinoneoida, as well as unravelling the relationships between sand dollars and their closest relatives. We then use this topology to trace the evolutionary history of echinoid body size through more than 270 million years, revealing a complex pattern of convergent evolution to stable peaks in macroevolutionary adaptive landscape. Our efforts show how combining phylogenomic and paleontological evidence offers new ways of exploring evolutionary forces operating across deep timescales.

Despite significant advances in invertebrate phylogenomics over the past decade, the higher-level phylogeny of Pycnogonida (sea spiders) remains elusive. Due to the inaccessibility of some small-bodied lineages, few phylogenetic studies have sampled all sea spider families. Previous efforts based on a handful of genes have yielded unstable tree topologies. Here, we inferred the relationships of 89 sea spider species using targeted capture of the mitochondrial genome, 56 conserved exons, 101 ultraconserved elements, and three nuclear ribosomal genes. We inferred molecular divergence times by integrating morphological data for fossil species to calibrate 15 nodes in the arthropod tree of life. This integration of data classes resolved the basal topology of sea spiders with high support. The enigmatic family Austrodecidae was resolved as the sister group to the remaining Pycnogonida and the small-bodied family Rhynchothoracidae as the sister group of the robust-bodied family Pycnogonidae. Molecular divergence time estimation recovered a basal divergence of crown group sea spiders in the Ordovician. Comparison of diversification dynamics with other marine invertebrate taxa that originated in the Paleozoic suggests that sea spiders and some crustacean groups exhibit resilience to mass extinction episodes, relative to mollusk and echinoderm lineages.

Nudibranch molluscs of the family Tritoniidae are widely used neuroscience model systems for understand the behavioural and genetic bases of learning and memory. However species identity and genus-level taxonomic assignment of the tritoniids remain contested. Herein we present a taxonomic review of the family Tritoniidae using integration of molecular phylogenetic analysis, morphological and biogeographical data. For the first time the identity of the model speciesTritonia tetraquetra(Pallas, 1788) andTritonia exsulansBergh, 1894 is confirmed.T.tetraquetradistributes across the large geographic and bathymetric distances in the North-Eastern (NE) and North-Western (NW) Pacific. In turn, at NE Pacific coasts the separate speciesT.exsulansis commonly occured. Thus, it reveals a misidentification ofT.tetraquetraandT.exsulansspecies in neuroscience applications. Presence of more hidden lineages within NW PacificT.tetraquetrais suggested. The long lasting confusion over identity of the species from the generaTritoniaandTochuinais resolved using molecular and morphological data. We also disprove a common indication about “edibleT.tetraquetra” at the Kuril Islands. It is shown thatTochuinapossesses specialized tritoniid features and also some characters of “arminacean nudibranchs”, such asDoridoxaandHeterodoris. Diagnoses for the families Doridoxidae and Heterodorididae are provided. Taxonomy of the genusDoridoxais clarified and molecular data for the genusHeterodorispresented for the first time. A taxonomic synopsis for the family Tritoniidae is provided. A new genus among tritoniid taxa is proposed. Importance of the ontogeny-based taxonomy is highlighted. The cases when apomorphic characters considerably modified in a crown group due to the paedomorphosis are revealed. Tracing of the character evolution is presented for secondary gills–a key external feature of the family Tritoniidae and traditional dendronotacean nudibranchs.

In the present study, a series of phylogenetic analyses of morphological, molecular, and combined morphological-molecular datasets were conducted to investigate the relationships of 23 extant and 44 fossil caniforme genera, in order to test the phylogenetic position of putative stem pinniped Puijila within a comprehensive evolutionary framework. With Canis as an outgroup, a Bayesian Inference analysis employing tip-dating of a combined molecular-morphological (i.e., Total Evidence) dataset recovered a topology in which musteloids are the sister group to a monophyletic pinniped clade, to the exclusion of ursids, and recovered Puijila and Potamotherium along the stem of Pinnipedia. A similar topology was recovered in a parsimony analysis of the same dataset. These results suggest the pinniped stem may be expanded to include additional fossil arctoid taxa, including Puijila, Potamotherium, and Kolponomos. The tip-dating analysis suggested a divergence time between pinnipeds and musteloids of ~45.16 million years ago (Ma), though a basal split between otarioids and phocids is not estimated to occur until ~26.52 Ma. These results provide further support for prolonged freshwater and nearshore phases in the evolution of pinnipeds, prior to the evolution of the extreme level of aquatic adaptation displayed by extant taxa. Ancestral character state reconstruction was used to investigate character evolution, to determine the frequency of reversals and parallelisms characterizing the three extant clades within Pinnipedia. Although the phylogenetic analyses did not directly provide any evidence of parallel evolution within the pinniped extant families, it is apparent from the inspection of previously-proposed pinniped synapomorphies, within the context of a molecular-based phylogenetic framework, that many traits shared between extant pinnipeds have arisen independently in the three clades. Notably, those traits relating to homodonty and limb-bone specialization for aquatic locomotion appear to have multiple origins within the crown group, as suggested by the retention of the plesiomorphic conditions in early-diverging fossil members of the three extant families. Thus, while the present analysis identifies a new suite of morphological synapomorphies for Pinnipedia, the frequency of reversals and other homoplasies within the clade limit their diagnostic value.

Featherwing beetles, or Ptiliidae (Coleoptera: Staphylinoidea), are characterized by their minute sized-body and peculiar hind wing structures, and include the smallest free-living insects. A half millimeter-long, unsexed adult featherwing beetle from Upper Cretaceous Burmese amber is here described as Kekveus jason gen. et sp. nov. Our observation recovered fine morphological details, such as the fringes of hind wings and pronotal grooves, from the Mesozoic Ptiliidae for the first time. Among Ptiliidae, this is the first nominal extinct genus, the first named representative from Burmese amber, the oldest named fossil, and the first named fossil from the Mesozoic Era. The new taxon is phylogenetically nested deep inside the crown group of Ptiliidae and is hypothesized to belong to the extant tribe Discheramocephalini, as either a member of, a sister to, the clade formed by the genera Skidmorella and Discheramocephalus. Our finding indicates the miniaturization in Ptiliidae can be reliably traced back to at least mid-Cretaceous.

The first comprehensive timetree is presented for phytoplasmas, a diverse group of obligate intracellular bacteria restricted to phloem sieve elements of vascular plants and tissues of their hemipteran insect vectors. Maximum likelihood-based phylogenetic analysis of DNA sequence data from the 16S rRNA and methionine aminopeptidase (map) genes yielded well resolved estimates of phylogenetic relationships among major phytoplasma lineages, 16Sr groups and known strains of phytoplasmas. Age estimates for divergences among two major lineages of Mollicutes based on a previous comprehensive bacterial timetree were used to calibrate an initial 16S timetree. A separate timetree was estimated based on the more rapidly-evolving map gene, with an internal calibration based on a recent divergence within two related 16Sr phytoplasma subgroups in group 16SrV thought to have been driven by the introduction of the North American leafhopper vector Scaphoideus titanus Ball into Europe during the early part of the 20th century. Combining the resulting divergence time estimates into a final 16S timetree suggests that evolutionary rates have remained relatively constant overall through the evolution of phytoplasmas and that the origin of this lineage, at ~641 million years ago (Ma), preceded the origin of land plants and hemipteran insects. Nevertheless, the crown group of phytoplasmas is estimated to have begun diversifying ~316 Ma, roughly coinciding with the origin of seed plants and Hemiptera. Some phytoplasma groups apparently associated with particular plant families or insect vector lineages generally arose more recently than their respective hosts and vectors, suggesting that vector-mediated host shifts have been an important mechanism in the evolutionary diversification of phytoplasmas. Further progress in understanding macroevolutionary patterns in phytoplasmas is hindered by large gaps in knowledge of the identity of competent vectors and lack of data on phytoplasma associations with non-economically important plants.

Preprint available on bioRxiv (https://doi.org/10.1101/826552) Abstract: Taxon sampling is a central aspect of phylogenetic study design, but it has received limited attention in the context of molecular dating and especially in the framework of total-evidence dating, a widely used dating approach that directly integrates molecular and morphological information from extant and fossil taxa. We here assess the impact of different outgroup sampling schemes on age estimates in a total-evidence dating analysis under the uniform tree prior. Our study group are Pimpliformes, a highly diverse, rapidly radiating group of parasitoid wasps of the family Ichneumonidae. We cover 201 extant and 79 fossil taxa, including the oldest fossils of the family from the Early Cretaceous and the first unequivocal representatives of extant subfamilies from the mid Paleogene. Based on newly compiled molecular data from ten nuclear genes and a morphological matrix that includes 222 characters, we show that age estimates become both older and less precise with the inclusion of more distant and more poorly sampled outgroups. In addition, we discover an artefact that might be detrimental for total-evidence dating: “bare-branch attraction”, namely high attachment probabilities of, especially, older fossils to terminal branches for which morphological data are missing. After restricting outgroup sampling and adding morphological data for the previously attracting, bare branches, we recover a Middle and Early Jurassic origin for Pimpliformes and Ichneumonidae, respectively. This first age estimate for the group not only suggests an older origin than previously thought, but also that diversification of the crown group happened before the Cretaceous-Paleogene boundary. Our case study demonstrates that in order to obtain robust age estimates, total-evidence dating studies need to be based on a thorough and balanced sampling of both extant and fossil taxa, with the aim of minimizing evolutionary rate heterogeneity and missing morphological Information.

PREMISE OF THE STUDY Equisetum is the sole living representative of Sphenopsida, a clade with impressive species richness, a long fossil history dating back to the Devonian, and obscure relationships with other living pteridophytes. Based on molecular data, the crown group age of Equisetum is mid-Paleogene, although fossils with possible crown synapomorphies appear in the Triassic. The most widely circulated hypothesis states that the lineage of Equisetum derives from calamitaceans, but no comprehensive phylogenetic studies support the claim. Using a combined approach, we provide a comprehensive phylogenetic analysis of Equisetales, with special emphasis on the origin of genus Equisetum. METHODS We performed parsimony phylogenetic analyses to address relationships of 43 equisetalean species (15 extant, 28 extinct) using a combination of morphological and molecular characters. KEY RESULTS We recovered Equisetaceae + Neocalamites as sister to Calamitaceae + a clade of Angaran and Gondwanan horsetails, with the four groups forming a clade that is sister to Archaeocalamitaceae. The estimated age for the Equisetum crown group is mid-Mesozoic. CONCLUSIONS Modern horsetails are not nested within calamitaceans; instead, both groups have explored independent evolutionary trajectories since the Carboniferous. Diverse fossil taxon sampling helps to shed light on the position and relationships of equisetalean lineages, of which only a tiny remnant is present within the extant flora. Understanding these relationships and early character configurations of ancient plant clades as Equisetales provide useful tests of hypotheses about overall phylogenetic relationships of euphyllophytes and foundations for future tests of molecular dates with paleontological data.

PREMISE OF THE STUDY Recent clarification of the distribution of Marattiales through time provides the impetus for "total evidence" phylogenetic analyses of a major fern clade with a rich fossil record. These analyses serve as empirical tests for results from systematic analyses of living species and also of the belief that relationships among living species accurately reflect the overall pattern of phylogeny for clades with an extensive fossil record and a large percentage of extinction. METHODS Species of living and fossil Marattiaceae are analyzed employing a "total evidence approach" via maximum parsimony. Analyses were conducted using TNT implemented through WinClada. KEY RESULTS Systematic analyses of living species and of living + extinct species provide roughly concordant topologies for living taxa. However, living species of Marattiales are only one component of a much larger clade with two major subclades. One consists of Psaroniaceae and extends through time to at least the Early Cretaceous. The other consists of Marattiaceae and includes all living species. Various analyses support the generic-level clades of living species from earlier analyses, but the arrangement of such clades varies from analysis to analysis. CONCLUSIONS Marattiales is a monophyletic group that is extremely common in late Paleozoic and early Mesozoic deposits, with a stem group Psaroniaceae and a crown group Marattiaceae. Because Marattiaceae represents only a small component of overall marattialean diversity, living species alone neither account for evolutionary changes within the clade over time, nor accurately reflect the overall pattern of marattialean fern phylogeny.

The origins, evolutionary history and diversification of the Australian butterfly fauna are poorly known and uncertain. Two competing hypotheses have been proposed to explain the occurrence of butterflies on this isolated continental landmass. The common view is that all Australian butterflies entered the continent relatively recently from the northern hemisphere via Southeast Asia and/or mainland New Guinea (i.e. northern dispersal origin hypothesis). The alternative view is that part or all of the Australian butterfly fauna ultimately evolved in remnant or Southern Gondwana when Australia was connected to South America through Antarctica (i.e. Southern Gondwanan origin hypothesis). However, robust phylogenies with strong support for monophyly are lacking for the majority of Australian endemic butterfly lineages, thereby precluding determination of their systematic relationships and hence their geographic origins. Here, we use molecular data to reconstruct phylogenetic relationships of the globally distributed butterfly subtribe Coenonymphina (Satyrinae: Satyrini). This group represents a major component of the butterfly fauna of the wider Australasian region, with 19 genera and 71 species endemic to the region. Dating estimates extrapolated from secondary calibration sources indicate that the subtribe arose c. 48 Ma (95% credibility interval, 52–42 Ma), and the crown group first diverged in the Eocene (c. 44 Ma, 95% credibility interval 51–37 Ma). Rapid speciation events subsequently followed around the Eocence–Oligocene boundary, resulting in a near-hard polytomy comprising short basal branches with nodes that are difficult to resolve. Based on strongly supported phylogenetic relationships and estimates of divergence times, we conclude that the group probably had its origin in the fragment of Southern Gondwana consisting of Australia, Antarctica and South America. However, we are unable to rule out the northern dispersal scenario, particularly as Coenonymphina are closely related to a set of predominantly Asian lineages. Dispersal and extinction events following the final break-up of Gondwana have played a pivotal role in shaping the extant distributions of the group. (Less)

Aim: To infer divergence times and historical biogeography of the cosmopolitan lycophyte family Lycopodiaceae. Location: Worldwide. Methods: We generated time‐calibrated phylogenies of the Lycopodiaceae based on six regions of chloroplast DNA using a node‐dating approach implemented in beast with eight fossil calibrations. To investigate effects of among‐lineage substitution rate heterogeneity on divergence time estimation, we compared the performance of two relaxed clock models: an uncorrelated lognormal clock model and a random local clock (RLC) model. The historical biogeography of the family was inferred using two Bayesian models implemented in BioGeoBEARS. Results: Divergence time estimates for major groups of the Lycopodiaceae obtained using the two substitution clock models differed substantially, and the RLC model was a better fit. The Lycopodiaceae crown group age is estimated to be late Devonian, and most deep divergence events date to the Carboniferous, with most extant species diversity accumulating during the Cenozoic. The timing of divergences of major clades in the Lycopodiaceae corresponds to the breakup of the Pangaean and Gondwanan supercontinents. Long‐distance dispersal events are relatively common, but generally do not appear to be followed by subsequent radiations. Main conclusions: Accounting for among‐lineage substitution rate heterogeneity improves divergence time estimates for the Lycopodiaceae. The family has a deep evolutionary history, and continent‐scale vicariance events in the Mesozoic appear to have been associated with major cladogenesis events, with long‐distance dispersal playing a relatively minor role.

The oldest definitive pinniped fossils date from approximately 30.6–23 million years ago (Ma) in the North Pacific. Pinniped monophyly is consistently supported; the group shares a common ancestry with arctoid carnivorans, either ursids or musteloids. Crown pinnipeds comprise the Otariidae (fur seals and sea lions), Odobenidae (walruses), and Phocidae (seals), with paraphyletic “enaliarctines” falling outside the crown group. The position of extinct Desmatophocidae is debated; they are considered to be closely related to both otariids and odobenids or, alternatively, to phocids. Both otariids and odobenids are known from the North Pacific, diverging approximately 19 Ma, with phocids originating in the North Atlantic or Paratethys region 19–14 Ma. Our understanding of pinniped paleobiology has been enriched by studies that incorporate anatomical and behavioral data into a phylogenetic framework. There is now evidence for sexual dimorphism in the earliest pinnipeds, heralding polygynous breeding systems, followed by increased body sizes, diving capabilities, and diverse feeding strategies in later-diverging phocid and otarioid lineages.

Chiliadenus is a small genus in the Inuleae (Asteraceae), consisting of ten species with allopatric distributions along the southern edge of the Mediterranean Sea. The different species have restricted areas of distribution, with only one being more widely distributed. The first molecular phylogenetic study of the genus with complete sampling, as well as a biogeographic analysis of the origin and biogeographic patterns leading to the current diversity of Chiliadenus is presented. Results confirm Chiliadenus as monophyletic and placed as sister to Dittrichia. The ancestor of Chiliadenus is dated to have diverged from that of Dittrichia around 5.45 Ma ago, coinciding with the Messinian salinity crisis, whereas the Chiliadenus crown group is dated to 2.29 Ma, around 3 million years later. Ancestral area reconstructions show the crown group to likely have originated in the area around Morocco and northwestern Algeria, which is also the area where the early divergences have occurred. Chiliadenus has then later diverged and dispersed over the Mediterranean to its current distribution. The evolution of the Chiliadenus crown group coincides with the onset of the Mediterranean climate, and its evolution may be connected to the subsequent climatic changes.

Apiosphaeria guaranitica, the causal agent of brown crust disease of several bignoniaceous hosts, among them Handroanthus and Tabebuia species, has been traditionally placed in Phyllachoraceae, based exclusively on morphological studies, without supporting molecular evidence. Here, we provide molecular data for the link between sexual and asexual states of the fungus and elucidate the phylogeny of A. guaranitica. The multilocus phylogenetic analyses employed sequences from the 18S subunit (18S), 28S subunit (28S), and nuclear internal transcribed spacers (ITS1-5.8S-ITS2 = ITS) of the nuc rDNA, second-largest subunit of RNA polymerase II (RPB2), and translation elongation factor 1-α (TEF1) genetic loci. Estimates of the divergence time of this lineage were supported by fossil calibration (FC) and secondary calibration (SC) strategies. Our results indicate a natural placement of Apiosphaeria within Diaporthaceae (Diaporthales), where it represents an ancient lineage of the crown group of Diaporthaceae, diverging during the late Paleocene at 61.15 (FC) and 60.63 (SC) million years ago. This divergence time estimate within Diaporthales is based on Spataporthe taylori, a diaporthaceous fossil.

Algorachelus peregrinus is the oldest representative of the crown group Pleurodira known in Laurasia. The type locality of this bothremydid is Algora, situated in central Spain, at levels deposited during the uppermost middle–lowermost upper Cenomanian. A new excavation was recently carried out in this town. As a result, abundant material of Algorachelus peregrinus has been found. Several complete shells, as well as numerous partial carapaces and plastra, are presented here. The abundance of remains allows the analysis of the general patterns by which some of the shells of this littoral form were partially or totally disarticulated. The analysis of these remains not only shows several pathologies but also improves the knowledge about the anatomy and intraspecific variability of Algorachelus peregrinus. This new information allows the revision of other Cenomanian forms of Bothremydidae, both from the Middle East and from North America. Thus, Algorachelus is identified in these regions, being represented in the early or middle Cenomanian of Palestine by the new combination Algorachelus parvus, and in the uppermost Cenomanian of Utah by the new combination Algorachelus tibert. Therefore, a relatively fast and widely distributed geographic dispersion event is recognized, corresponding to the oldest dispersal event so far identified for a lineage of Pleurodira from Gondwana to Laurasia.

A new species of the Marattialean fern frond Rothwellopteris is proposed for specimens preserving both morphology and anatomy from the Wuchiapingian–Changshingian (late Permian) aged Xuanwei Formation of South China. Fronds are at least bipinnate and bear falcate, pecopteroid pinnules with a thin lamina. The penultimate pinna rachis has a polycyclic vascular system with endarch primary xylem and longer vascular bundles located abaxially in a single radial row, and shorter vascular bundles adaxially arranged in two radial rows, with a vascular bundle sheath of sclerenchyma cells. Ground tissue comprises parenchyma cells and numerous secretory ducts. Features of the new species suggest that it was adapted for mesomorphy, while the previously recognized species Rothwellopteris pecopteroides was adapted for xeromorphy or was a sun-leaf. Like R. pecopteroides, the new species displays a mosaic of morphological and anatomical features characteristic of both the Paleozoic family Psaroniaceae and the extant family Marattiaceae; we interpret it to be a member of the evolutionary stem-group from which stratigraphically younger Marattiaceae evolved. Our findings show that the stem- to crown-group transition within Marattiales commenced before the end of the Permian and that a greater diversity of transitional forms can now be recognized from the late Permian and include the genera Compsopteris, Rothwellopteris, Tietea and Tuvichapteris.

Balaenopteroidae is a diverse superfamily within the mysticetes that consists of numerous recent fossil species. The publication of new taxa has increased, which improves the knowledge about this group. In this work, we name and describe Kennedycetus pericorum from the Late Miocene deposits of the Trinidad Formation in Baja California Sur, Mexico. Our phylogenetic results indicate that this is a step taxon, far from the balaenopterid crown group, that is closely related to Norrisanima miocaena from the Monterey Formation, California. The associated fauna of K. pericorum includes the internal molds of different mollusks attributable to Glycymeridae, Semelidae, and Lucinidae, which indicate a coastal environment of warm waters.

Cololejeuneinae is the largest subtribe of the liverwort family Lejeuneaceae and the crown group of the family. The species often grow on the surfaces of living leaves and twigs or in swiftly running waters and many of them possess neotenic gametophytes, considered an adaptation to these extreme habitats. We describe a new, epiphyllous genus and species in Cololejeuneinae from the Western Cordillera of Ecuador, Reinerantha foliicola Gradst. & R.L.Zhu. Analysis of combined molecular datasets with parsimony, maximum likelihood and Bayesian methods revealed Reinerantha in a sister relationship to the large pantropical genus Diplasiolejeunea. Reinerantha differs from Diplasiolejeunea in leaf segmentation, epidermal wall thickening, stylus, male bracts and perianth keels, and is morphologically more similar to the genus Tuyamaella. In the molecular analysis, however, Reinerantha and Tuyamaella were resolved in different clades and not closely related. Tuyamaella is shown to be restricted to Asia; the report of Tuyamaella from South America (Peru) was erroneous and a case of mislabeling.

Hybridization and genome doubling (allopolyploidy) have led to evolutionary novelties as well as to the origin of new clades and species. Despite the importance of allopolyploidization, the dynamics of post-polyploid diploidization (PPD) at the genome level has been only sparsely studied. The Microlepidieae (MICR) is a crucifer tribe of 17 genera and c. 56 species endemic to Australia and New Zealand. Our phylogenetic and cytogenomic analyses revealed that MICR originated via an inter-tribal hybridization between ancestors of Crucihimalayeae (n = 8; maternal genome) and Smelowskieae (n = 7; paternal genome), both native to the Northern Hemisphere. The reconstructed ancestral allopolyploid genome (n = 15) originated probably in north-eastern Asia or western North America during the Late Miocene (c. 10.6 - 7 million years ago) and reached the Australian mainland via long-distance dispersal. In Australia, the allotetraploid genome diverged into at least three main subclades exhibiting different levels of PPD and diversity: 1.25-fold descending dysploidy (DD) of n = 15 → n = 12 (autopolyploidy → 24) in perennial Arabidella (3 species), 1.5-fold DD of n = 15 → n = 10 in the perennial Pachycladon (11 spp.), and 2.1 to 3.75-fold DD of n = 15 → n = 7 - 4 in the largely annual crown group genera (42 spp. in 15 genera). These results are among the first to demonstrate multi-speed genome evolution in taxa descending from a common allopolyploid ancestor. It is suggested that clade-specific PPD can operate at different rates and efficacies, and can be tentatively linked to life histories and the extent of taxonomic diversity. This article is protected by copyright. All rights reserved.