The oldest known fur seal

The poorly known fossil record of fur seals and sea lions (Otariidae) does not reflect their current diversity and widespread abundance. This limited fossil record contrasts with the more complete fossil records of other pinnipeds such as walruses (Odobenidae). The oldest known otariids appear 5–6 Ma after the earliest odobenids, and the remarkably derived craniodental morphology of otariids offers few clues to their early evolutionary history and phylogenetic affinities among pinnipeds. We report a new otariid, Eotaria crypta, from the lower middle Miocene ‘Topanga’ Formation (15–17.1 Ma) of southern California, represented by a partial mandible with well-preserved dentition. Eotaria crypta is geochronologically intermediate between ‘enaliarctine’ stem pinnipedimorphs (16.6–27 Ma) and previously described otariid fossils (7.3–12.5 Ma), as well as morphologically intermediate by retaining an M₂ and a reduced M₁ metaconid cusp and lacking P_2–4 metaconid cusps. Eotaria crypta eliminates the otariid ghost lineage and confirms that otariids evolved from an ‘enaliarctine’-like ancestor.     

The diversification of the northern temperate woody flora – A case study of the Elm family (Ulmaceae) based on phylogenomic and paleobotanical evidence

Ulmaceae is a woody family widespread in northern temperate forests. Despite the ecological importance of this family, its phylogeny and biogeographic history are poorly understood. In this study, we reconstruct phylogenetic relationships within the family and infer spatio-temporal diversification patterns based on chloroplast genome (complete cpDNA) and nuclear ribosomal DNA sequences (nrDNA). The seven Ulmaceae genera are resolved in two main clades (temperate vs. tropical) by both cpDNA and nrDNA sequences. The temperate clade includes four genera, Hemiptelea, Zelkova, Planera, and Ulmus. The relationships among Planera and other genera are controversial because of inconsistent topologies between plastid and nuclear data. The tropical clade includes three genera ((Ampelocera, Phyllostylon), Holoptelea). Molecular dating and diversification analyses show that Ulmaceae originated in the Early Cretaceous (ca. 110–125 Ma) with the main lineages establishing from the Late Cretaceous to the early Eocene. The diversification rate slowed during the middle to the late Paleogene (ca. 23–45 Ma), followed by a rapid diversification of the East Asian temperate group in the Neogene, congruent with a global cooling event. The ancestral state optimization analysis suggests an East Asian origin of the temperate Ulmaceae clade during the Paleocene, which is consistent with the fossil record. Both phylogenomic and fossil evidence support East Asia as a center of origin and diversification for the temperate woody lineages.     

Dating the origin of the major lineages of Branchiopoda

Despite the well-established phylogeny and good fossil record of branchiopods, a consistent macro-evolutionary timescale for the group remains elusive. This study focuses on the early branchiopod divergence dates where fossil record is extremely fragmentary or missing. On the basis of a large genomic dataset and carefully evaluated fossil calibration points, we assess the quality of the branchiopod fossil record by calibrating the tree against well-established first occurrences, providing paleontological estimates of divergence times and completeness of their fossil record. The maximum age constraints were set using a quantitative approach of Marshall (2008). We tested the alternative placements of Yicaris and Wujicaris in the referred arthropod tree via the likelihood checkpoints method. Divergence dates were calculated using Bayesian relaxed molecular clock and penalized likelihood methods. Our results show that the stem group of Branchiopoda is rooted in the late Neoproterozoic (563 ± 7 Ma); the crown-Branchiopoda diverged during middle Cambrian to Early Ordovician (478–512 Ma), likely representing the origin of the freshwater biota; the Phyllopoda clade diverged during Ordovician (448–480 Ma) and Diplostraca during Late Ordovician to early Silurian (430–457 Ma). By evaluating the congruence between the observed times of appearance of clade in the fossil record and the results derived from molecular data, we found that the uncorrelated rate model gave more congruent results for shallower divergence events whereas the auto-correlated rate model gives more congruent results for deeper events.     

Systematics and evolution of the Pan‐Alcidae (Aves,Charadriiformes)

Puffins, auks and their allies in the wing‐propelled diving seabird clade Pan‐Alcidae (Charadriiformes) have been proposed to be key pelagic indicators of faunal shifts in Northern Hemisphere oceans. However, most previous phylogenetic analyses of the clade have focused only on the 23 extant alcid species. Here we undertake a combined phylogenetic analysis of all previously published molecular sequence data (～ 12 kb) and morphological data (n = 353 characters) with dense species level sampling that also includes 28 extinct taxa. We present a new estimate of the patterns of diversification in the clade based on divergence time estimates that include a previously vetted set of twelve fossil calibrations. The resultant time trees are also used in the evaluation of previously hypothesized paleoclimatic drivers of pan‐alcid evolution. Our divergence dating results estimate the split of Alcidae from its sister taxon Stercorariidae during the late Eocene (～ 35 Ma), an evolutionary hypothesis for clade origination that agrees with the fossil record and that does not require the inference of extensive ghost lineages. The extant dovekie Alle alle is identified as the sole extant member of a clade including four extinct Miocene species. Furthermore, whereas an Uria + Alle clade has been previously recovered from molecular analyses, the extinct diversity of closely related Miocepphus species yields morphological support for this clade. Our results suggest that extant alcid diversity is a function of Miocene diversification and differential extinction at the Pliocene–Pleistocene boundary. The relative timing of the Middle Miocene climatic optimum and the Pliocene–Pleistocene climatic transition and major diversification and extinction events in Pan‐Alcidae, respectively, are consistent with a potential link between major paleoclimatic events and pan‐alcid cladogenesis.     

Divergence Times and the Evolutionary Radiation of New World Monkeys (Platyrrhini,Primates): An Analysis of Fossil and Molecular Data

The estimation of phylogenetic relationships and divergence times among a group of organisms is a fundamental first step toward understanding its biological diversification. The time of the most recent or last common ancestor (LCA) of extant platyrrhines is one of the most controversial among scholars of primate evolution. Here we use two molecular based approaches to date the initial divergence of the platyrrhine clade, Bayesian estimations under a relaxed-clock model and substitution rate plus generation time and body size, employing the fossil record and genome datasets. We also explore the robustness of our estimations with respect to changes in topology, fossil constraints and substitution rate, and discuss the implications of our findings for understanding the platyrrhine radiation. Our results suggest that fossil constraints, topology and substitution rate have an important influence on our divergence time estimates. Bayesian estimates using conservative but realistic fossil constraints suggest that the LCA of extant platyrrhines existed at ca. 29 Ma, with the 95% confidence limit for the node ranging from 27–31 Ma. The LCA of extant platyrrhine monkeys based on substitution rate corrected by generation time and body size was established between 21–29 Ma. The estimates based on the two approaches used in this study recalibrate the ages of the major platyrrhine clades and corroborate the hypothesis that they constitute very old lineages. These results can help reconcile several controversial points concerning the affinities of key early Miocene fossils that have arisen among paleontologists and molecular systematists. However, they cannot resolve the controversy of whether these fossil species truly belong to the extant lineages or to a stem platyrrhine clade. That question can only be resolved by morphology. Finally, we show that the use of different approaches and well supported fossil information gives a more robust divergence time estimate of a clade.     

The fossil record of Limopsis (Bivalvia: Limopsidae) in Antarctica and the southern high latitudes

Abstract: Limopsis is one of the most speciose and widespread bivalve genera in the Southern Ocean at the present day. However, the fossil record of the genus is poorly known from the southern high latitudes. Here, we review the fossil record in this region to help understand the evolutionary origins of the genus. Limopsis infericola sp. nov. and additional specimens of a previously described species are added to the fossil record of Antarctica. The globally distributed limopsid clade had its earliest occurrences in the Early Cretaceous of Europe and New Zealand, then radiated during the Late Cretaceous (Maastrichtian, 70.6–65.5 Ma). Fossil evidence shows that the genus underwent a second, Cenozoic, radiation related to the isolation of Antarctica and the onset of cooling in the southern hemisphere. The genus has persisted in Antarctica for the last 50 myr, adapting to extreme changes in the environmental conditions, including surviving the last glacial maximum in marine refugia.     

Revisiting taxonomy, morphological evolution, and fossil calibration strategies in Chloranthaceae

Chtoranthaceae is one of the earliest diverging angiosperm families and is comprised of approximately 75 species in four genera (Chloranthus,Sarcandra,Ascarina,and Hedyosmurn).This family has received considerable attention because of its seemingly primitive morphology,disjunct tropical distribution in Asia and America,and extensive fossil record from the Early Cretaceous.In the present study,we reconstructed the phylogeny of Chloranthaceae based on a combined dataset of three plastid DNA regions and 56 species.We then estimated divergence times in the family using two relaxed molecular clock methods (BEAST and penalized likelihood).We focused on testing the influence of fossil taxa in calibrating the molecular phylogeny,and on assessing the current taxonomy of the family in light of the phylogenetic results.Our results indicate that most intrageneric divisions within Ascarina and Hedyosmum are not monophyletic.The results from the dating analysis suggest that the Hedyosmum-like fossil Asteropollis represents a stem lineage of Hedyosmum,as has been suggested previously from morphological analyses.In contrast,our results indicate that the Chloranthus-like fossil Chloranthistemon,previously suggested on morphological grounds to be a stem relative of Chloranthus,may,instead,belong to the branch leading to the clade Chloranthus + Sarcandra.The median crown ages of Chloranthus,Sarcandra,Ascarina,and Hedyosmum estimated in the BEAST analysis were 26.3,9.5,31.0 and 45.8 million years ago (Ma),respectively,whereas the divergence between Chloranthus and Sarcandra,the splitting of Ascarina with the former two genera,and Hedyosmum separating from the three genera were estimated to 63.8,95.7 and 111.1 Ma.The present study sheds further light on the temporal evolution of Chloranthaceae and exemplifies how molecular dating analyses may be used to explore alternative phylogenetic placements of fossil taxa.     

A time-calibrated multi-gene phylogeny of the diatom genus Pinnularia

Pinnularia is an ecologically important and species-rich genus of freshwater diatoms (Bacillariophyceae) showing considerable variation in frustule morphology. Interspecific evolutionary relationships were inferred for 36 Pinnularia taxa using a five-locus dataset. A range of fossil taxa, including newly discovered Middle Eocene forms of Pinnularia, was used to calibrate a relaxed molecular clock analysis and investigate temporal aspects of the genus' diversification. The multi-gene approach resulted in a well-resolved phylogeny of three major clades and several subclades that were frequently, but not universally, delimited by valve morphology. The genus Caloneis was not recovered as monophyletic, confirming that, as currently delimited, this genus is not evolutionarily meaningful and should be merged with Pinnularia. The Pinnularia-Caloneis complex is estimated to have diverged between the Upper Cretaceous and the early Eocene, implying a ghost range of at least 10 million year (Ma) in the fossil record.     

Three extant genera of freshwater thalassiosiroid diatoms from Middle Eocene sediments in northern Canada

The evolutionary history of diatoms is only constrained partially by the fossil record. The timing of several key events, such as initial colonization of freshwater habitats by marine taxa, remains poorly resolved. Numerous specimens of the genera Cyclotella, Discostella, and Puncticulata (Ochrophyta: Thalassiosirales) have been recovered in Middle Eocene lacustrine sediments from the Giraffe Pipe locality in the Northwest Territories, Canada. These diatoms extend the fossil record of the family Stephanodiscaceae to at least 40 million years before present (Ma) and thus provide a new evolutionary milepost for the thalassiosiroid diatoms, an important clade of centric diatoms with global representation in both marine and freshwater environments. The quality of the fossil material enables detailed investigations of areolae, fultoportulae, and rimoportulae, revealing direct morphological affinities with a number of extant taxa. These observations extend the antiquity of several characters of phylogenetic importance within the thalassiosiroid diatoms, including the fultoportula, and imply that the entire lineage is considerably older than prior constraints from the fossil record, as suggested independently by several recent molecular phylogenies.     

生物古地理、分子分异时间及化石纪录:古中华貂在内蒙古阿巴嘎旗晚中新世宝格达乌拉组的新材料(英文)

北美现生的潘氏貂Martes pennanti(荷兰殖民区俗名fisher,北美印地安土著俗名pe-kan)传统被归入貂属(Martes)。20世纪70年代初Elaine Anderson首先认识到北美fisher的祖先可以追溯到中国山西保德晚中新世的古中华貂M.palaeosinensis。它们的共同特征是上裂齿唇侧具有一个孤立的小齿根。因此在众多的化石和现生貂中,这些具有多一个小齿根的种类可以归入它们自己的一个分支,并重新启用已建立的Pekania属来区别于其他种类。近年来新兴的分子系统研究也趋于把北美的fisher与中、南美的Eira及北半球的狼獾(Gulo)并列,甚至指示它可能是狼獾的姊妹群。因此把fisher独立归入Pekania属中可能是未来的方向。2009年我们在内蒙古阿巴嘎旗宝格达乌拉组的晚中新世地层中发现了一个古中华貂的上颌。这是该种在内蒙古的首次发现,借此机会对目前的分子系统关系、分异年代及化石记录进行综合探讨。虽然保德和宝格达乌拉的古中华貂在时代上与分子钟估算的5～8Ma比较一致,狼獾的化石记录却远比古中华貂的老(可以追到11Ma左右)。如果狼獾确实是fisher的姊妹群或与它并系的话,那么fisher的分异时间也可能更早。     

Combining geometric morphometrics,molecular phylogeny,and micropaleontology to assess evolutionary patterns in Mallomonas (Synurophyceae: Heterokontophyta)

Synurophytes, also known as scaled chrysophytes, are ecologically important algae that produce an array of siliceous structures upon which their taxonomy is based. Despite occupying a key position within the photosynthetic heterokonts, the evolutionary history of synurophytes remains poorly constrained. Here, modern and Middle Eocene siliceous scales of the morphotaxon Mallomonas insignis are used as a model to investigate synurophyte evolutionary patterns. Structural details of scale morphology were examined comparatively with scanning electron microscopy and scored for geometric morphometric analyses to assess the stability of shape characters. Although consistent size differences exist (modern scales are larger than Eocene counterparts), the populations cannot be differentiated on the basis of shape or microstructural detail, implying considerable evolutionary stasis in scale morphology. A time‐calibrated relaxed molecular clock analysis using a three‐gene concatenated data set (27 strains) suggests that the M. insignis lineage predates the available fossil record, having diverged from closest congeneric taxa in the Cretaceous (≥94 Ma). However, the molecular analysis also implies that considerable genetic variability is present within several morphotaxa of Mallomonas, implying that substantial genetic variability has arisen despite the retention of uniform scale morphologies, and resulting in the widespread occurrence of cryptic taxa. Results from the synurophyte lineage are consistent with the notion of protracted ghost ranges (>10 Ma) implied by the molecular phylogenies of other algal groups, together pointing to the paucity of the fossil record of these organisms on these timescales.     

An Early Pliocene Hipparionine Horse from the Canadian Arctic

A partial skull of a juvenile hipparionine equid from Ellesmere Island, Canada, is the northernmost fossil record of a horse (78° 33' N). Biostratigraphical analysis of the associated fossil biota suggests an age of 3.5 to 4 Ma (early Pliocene). Preserved facial characteristics of the equid include a very reduced preorbital fossa located posterior to the infraorbital foramen. The deciduous premolars have low crown heights, complex fossette plications, multiple pli caballins, and oval, isolated protocones. The teeth are quite large, corresponding to an adult with a tooth row length of c. 150 mm. This combination of facial and dental characteristics and large size is not observed in any contemporaneous North American hipparionine, but is instead found in some Asiatic hipparionines, most notably Plesiohipparion . If the resemblance is not a result of convergence, then this represents the first record of an Old World hipparionine dispersing to North America. Alternatively, the specimen may represent a hitherto unknown, high-latitude hipparionine clade.     

Fossil ghost ranges are most common in some of the oldest and some of the youngest strata

Biologists routinely compare inferences about the order of evolutionary branching (phylogeny) with the order in which groups appear in the fossil record (stratigraphy). Where they conflict, ghost ranges are inferred: intervals of geological time where a fossil lineage should exist, but for which there is no direct evidence. The presence of very numerous and/or extensive ghost ranges is often believed to imply spurious phylogenies or a misleadingly patchy fossil record, or both. It has usually been assumed that the frequency of ghost ranges should increase with the age of rocks. Previous studies measuring ghost ranges for whole trees in just a small number of temporal bins have found no significant increase with antiquity. This study uses a much higher resolution approach to investigate the gappiness implied by 1,000 animal and plant cladograms over 77 series and stages of the Phanerozoic. It demonstrates that ghost ranges are indeed relatively common in some of the oldest strata. Surprisingly, however, ghost ranges are also relatively common in some of the youngest, fossil-rich rocks. This pattern results from the interplay between several complex factors and is not a simple function of the completeness of the fossil record. The Early Palaeozoic record is likely to be less organismically and stratigraphically complete, and its fossils -- many of which are invertebrates-may be more difficult to analyse cladistically. The Late Cenozoic is subject to the pull of the Recent, but this accounts only partially for the increased gappiness in the younger strata.     

Discriminating signal from noise in the fossil record of early vertebrates reveals cryptic evolutionary history

The fossil record of early vertebrates has been influential in elucidating the evolutionary assembly of the gnathostome bodyplan. Understanding of the timing and tempo of vertebrate innovations remains, however, mired in a literal reading of the fossil record. Early jawless vertebrates (ostracoderms) exhibit restriction to shallow-water environments. The distribution of their stratigraphic occurrences therefore reflects not only flux in diversity, but also secular variation in facies representation of the rock record. Using stratigraphic, phylogenetic and palaeoenvironmental data, we assessed the veracity of the fossil records of the jawless relatives of jawed vertebrates (Osteostraci, Galeaspida, Thelodonti, Heterostraci). Non-random models of fossil recovery potential using Palaeozoic sea-level changes were used to calculate confidence intervals of clade origins. These intervals extend the timescale for possible origins into the Upper Ordovician; these estimates ameliorate the long ghost lineages inferred for Osteostraci, Galeaspida and Heterostraci, given their known stratigraphic occurrences and stem–gnathostome phylogeny. Diversity changes through the Silurian and Devonian were found to lie within the expected limits predicted from estimates of fossil record quality indicating that it is geological, rather than biological factors, that are responsible for shifts in diversity. Environmental restriction also appears to belie ostracoderm extinction and demise rather than competition with jawed vertebrates.     

Phylogenetic relationships and divergence dates of softshell turtles (Testudines: Trionychidae) inferred from complete mitochondrial genomes

The softshell turtles (Trionychidae) are one of the most widely distributed reptile groups in the world, and fossils have been found on all continents except Antarctica. The phylogenetic relationships among members of this group have been previously studied; however, disagreements regarding its taxonomy, its phylogeography and divergence times are still poorly understood as well. Here, we present a comprehensive mitogenomic study of softshell turtles. We sequenced the complete mitochondrial genomes of 10 softshell turtles, in addition to the GenBank sequence of Dogania subplana, Lissemys punctata, Trionyx triunguis, which cover all extant genera within Trionychidae except for Cyclanorbis and Cycloderma. These data were combined with other mitogenomes of turtles for phylogenetic analyses. Divergence time calibration and ancestral reconstruction were calculated using BEAST and RASP software, respectively. Our phylogenetic analyses indicate that Trionychidae is the sister taxon of Carettochelyidae, and support the monophyly of Trionychinae and Cyclanorbinae, which is consistent with morphological data and molecular analysis. Our phylogenetic analyses have established a sister taxon relationship between the Asian Rafetus and the Asian Palea + Pelodiscus + Dogania + Nilssonia + Amyda, whereas a previous study grouped the Asian Rafetus with the American Apalone. The results of divergence time estimates and area ancestral reconstruction show that extant Trionychidae originated in Asia at around 108 million years ago (MA), and radiations mainly occurred during two warm periods, namely Late Cretaceous–Early Eocene and Oligocene. By combining the estimated divergence time and the reconstructed ancestral area of softshell turtles, we determined that the dispersal of softshell turtles out of Asia may have taken three routes. Furthermore, the times of dispersal seem to be in agreement with the time of the India–Asia collision and opening of the Bering Strait, which provide evidence for the accuracy of our estimation of divergence time. Overall, the mitogenomes of this group were used to explore the origin and dispersal route of Trionychidae and have provided new insights on the evolution of this group.     

A new Early Permian reptile and its significance in early diapsid evolution

The initial stages of evolution of Diapsida (the large clade that includes not only snakes, lizards, crocodiles and birds, but also dinosaurs and numerous other extinct taxa) is clouded by an exceedingly poor Palaeozoic fossil record. Previous studies had indicated a 38 Myr gap between the first appearance of the oldest diapsid clade (Araeoscelidia), ca 304 million years ago (Ma), and that of its sister group in the Middle Permian (ca 266 Ma). Two new reptile skulls from the Richards Spur locality, Lower Permian of Oklahoma, represent a new diapsid reptile: Orovenator mayorum n. gen. et sp. A phylogenetic analysis identifies O. mayorum as the oldest and most basal member of the araeoscelidian sister group. As Richards Spur has recently been dated to 289 Ma, the new diapsid neatly spans the above gap by appearing 15 Myr after the origin of Diapsida. The presence of O. mayorum at Richards Spur, which records a diverse upland fauna, suggests that initial stages in the evolution of non-araeoscelidian diapsids may have been tied to upland environments. This hypothesis is consonant with the overall scant record for non-araeoscelidian diapsids during the Permian Period, when the well-known terrestrial vertebrate communities are preserved almost exclusively in lowland deltaic, flood plain and lacustrine sedimentary rocks.     

Molecular phylogenetics and evolution of turtles

Turtles are one of Earth's most instantly recognizable life forms, distinguished for over 200 million years in the fossil record. Even so, key nodes in the phylogeny of turtles remain uncertain. To address this issue, we sequenced >90% of the nuclear recombination activase gene 1 (RAG-1) for 24 species representing all modern turtle families. RAG-1 exhibited negligible saturation and base composition bias, and extensive base composition homogeneity. Most of the relationships suggested by prior phylogenetic analyses were also supported by RAG-1 and, for at least two critical nodes, with a much higher level of support. RAG-1 also indicates that the enigmatic Platysternidae and Chelydridae, often considered sister taxa based on morphological evidence, are not closely related, although their precise phylogenetic placement in the turtle tree is still unresolved. Although RAG-1 is phylogenetically informative, our research revealed fundamental conflicts among analytical methods for estimating phylogenetic hypotheses. Maximum parsimony analyses of RAG-1 alone and in combination with two mitochondrial genes suggest the earliest phylogenetic splits separating into three basal branches, the pig-nosed turtles (Carettochelyidae), the softshell turtles (Trionychidae), and a clade comprising all remaining extant turtles. Maximum likelihood and Bayesian analyses group Carettochelyidae and Trionychidae (=Trionychoidae) in their more traditional location as the sister taxon to all other hidden-necked turtles, collectively forming the Cryptodira. Our research highlights the utility of molecular data in identifying issues of character homology in morphological datasets, while shedding valuable light on the biodiversity of a globally imperiled taxon.     

Detarieae sensu lato (Fabaceae) from the Late Oligocene (27.23 Ma) Guang River flora of north‐western Ethiopia

New species of caesalpinioid legumes, Cynometra sensu lato and Afzelia, are described from the Late Oligocene (27.23 Ma) Guang River flora in north‐western Ethiopia. Both taxa show leaf characteristics that are shared with extant species in the Guineo‐Congolian, Sudanian and/or Zambezian regions of Africa today. The presence of these two species in Ethiopia during the Palaeogene provides further evidence of the importance of the legume tribe Detarieae in northern and north‐eastern Africa throughout much of the Cenozoic, even although the clade is poorly represented in these regions today. The fossil record documents a significant palaeogeographical and evolutionary history of Detarieae in Africa, especially compared with that of Europe and Anatolia. Based on this evidence, it is unlikely that significant diversification of extant African Detarieae took place on the Eurasian landmass. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163 , 44–54.     

Cratonia cotyledon gen. et sp. nov: a unique Cretaceous seedling related to Welwitschia

The fossil history of most extant seed plant groups is relatively well documented. Cycads, conifers and Ginkgo all have an extensive fossil record, and the understanding of early angiosperm diversity is increasing. The Gnetales are an exception. Few macrofossils have been described, and character evolution within the group is poorly known. Cratonia cotyledon is a new gnetalean fossil from the Early Cretaceous Crato Formation of Brazil. This well-preserved seedling consists of two cotyledons, a feeder and a root. The leaf surface shows polygonal epidermal cells and apparently paracytic or actinocytic stomata. The cotyledons have a very specific venation pattern, shared only by Cratonia and Welwitschia, with parallel primary veins and secondary veins fusing to form inverted 'Y's between the main veins. Based on the 'Y'-venation and the presence of a feeder, we assign Cratonia to the Gnetum-Welwitschia clade. Fossil seedlings are unusual and this complete specimen with unambiguously welwitschioid characters is spectacular. Cratonia indicates that the evolutionary split between Gnetum and Welwitschia had occurred in the Early Cretaceous. Further, the close relationship between a West African plant and an east South American Early Cretaceous fossil is consistent with a major geological event: the rifting of the Gondwana continent.