A giant stem-group chaetognath.

Chaetognaths, with their characteristic grasping spines, are the oldest known pelagic predators, found in the lowest Cambrian (Terreneuvian). Here, we describe a large stem chaetognath, Timorebestia koprii gen. et sp. nov., from the lower Cambrian Sirius Passet Lagerstätte, which exhibits lateral and caudal fins, a distinct head region with long antennae and a jaw apparatus similar to Amiskwia sagittiformis. Amiskwia has previously been interpreted as a total-group chaetognathiferan, as either a stem-chaetognath or gnathostomulid. We show that T. koprii shares a ventral ganglion with chaetognaths to the exclusion of other animal groups, firmly placing these fossils on the chaetognath stem. The large size (up to 30 cm) and gut contents in T. koprii suggest that early chaetognaths occupied a higher trophic position in pelagic food chains than today.

Putting heads together.

Cambrian fossils reveal ancestry of the segmented brain in arthropods.

Exceptional soft tissue preservation reveals a cnidarian affinity for a Cambrian phosphatic tubicolous enigma.

Exoskeletal dwelling tubes are widespread among extant animals and early fossil assemblages. Exceptional fossils from the Cambrian reveal independent origins of tube dwelling by several clades including cnidarians, lophophorates, annelids, scalidophorans, panarthropods and ambulacrarians. However, most fossil tubes lack preservation of soft parts, making it difficult to understand their affinities and evolutionary significance. Gangtoucunia aspera (Wulongqing Formation, Cambrian Stage 4) was an annulated, gradually expanding phosphatic tube, with occasional attachments of multiple, smaller juveniles and has previously been interpreted as the dwelling tube of a 'worm' (e.g. a scalidophoran), lophophorate or problematicum. Here, we report the first soft tissues from Gangtoucunia that reveal a smooth body with circumoral tentacles and a blind, spacious gut that is partitioned by septa. This is consistent with cnidarian polyps and phylogenetic analysis resolves Gangtoucunia as a total group medusozoan. The tube of Gangtoucunia is phenotypically similar to problematic annulated tubular fossils (e.g. Sphenothallus, Byronia, hyolithelminths), which have been compared to both cnidarians and annelids, and are among the oldest assemblages of skeletal fossils. The cnidarian characters of G. aspera suggest that these early tubular taxa are best interpreted as cnidarians rather than sessile bilaterians in the absence of contrary soft tissue evidence.

A Cambrian tommotiid preserving soft tissues reveals the metameric ancestry of lophophorates.

Among extant animals, Lophotrochozoa accounts for the majority of phyla.1 This bilaterian clade radiated rapidly during the Cambrian explosion, obfuscating its phylogenetic relationships and rendering many aspects of its early evolution uncertain. Many early lophotrochozoans are known only from isolated skeletal microfossils, "small shelly fossils," often derived from larger animals with complex multi-element skeletons.2 The discovery of articulated fossils has revealed surprising insights into the animals from which these skeletal pieces were derived, such as paired shells in the mollusc Halkieria.3 Tommotiids are a key group of phosphatic early skeletal fossils that first appear in the late early Cambrian.4,5 Although their affinities were previously obscure, discoveries of partial scleritomes and investigations of growth and microstructure6 provide links with Brachiopoda7,8 and Phoronida,9 two of the lophophorate phyla. By contrast, the body plan of camenellan tommotiids remains a palaeontological mystery, with hypothetical reconstructions representing motile, benthic, dorsally armored worms.4,10 Here, we describe an articulated camenellan (Wufengella bengtsoni gen. et sp. nov.) from the Cambrian Chengjiang Biota, China, revealing the morphology of the scleritome and the first soft tissues from an adult tommotiid. Wufengella carries two dorsal rows of sclerites in a highly asymmetric arrangement, flanked by smaller, cap-shaped sclerites. The scleritome was fringed by iterated fascicles of chaetae and two layers of flattened lobes. Phylogenetic analysis confirms that camenellans occupy a deep branch in lophophorate phylogeny, prior to the acquisition of a sessile lifestyle. Wufengella reveals direct evidence for a metameric body plan reminiscent of annelids early in the evolutionary history of lophophorates.11,12.

Cambrian comb jellies from Utah illuminate the early evolution of nervous and sensory systems in ctenophores.

Ctenophores are a group of predatory macroinvertebrates whose controversial phylogenetic position has prompted several competing hypotheses regarding the evolution of animal organ systems. Although ctenophores date back at least to the Cambrian, they have a poor fossil record due to their gelatinous bodies. Here, we describe two ctenophore species from the Cambrian of Utah, which illuminate the early evolution of nervous and sensory features in the phylum. Thalassostaphylos elegans has 16 comb rows, an oral skirt, and an apical organ with polar fields. Ctenorhabdotus campanelliformis has 24 comb rows, an oral skirt, an apical organ enclosed by a capsule and neurological tissues preserved as carbonaceous films. These are concentrated around the apical organ and ciliated furrows, which connect to a circumoral nerve ring via longitudinal axons. C. campanelliformis deviates from the neuroanatomy of living ctenophores and demonstrates a substantial complexity in the nervous system of Cambrian ctenophores.

Evolution: No extinction? No way!

The fossil record reveals rampant extinction. However, analyses of time-calibrated molecular phylogenies often find no extinction at all. A new paper shows that estimates of zero extinction are entirely incorrect and are caused by limitations of analysing phylogenies that sample only living species.

Fossils improve phylogenetic analyses of morphological characters.

Fossils provide our only direct window into evolutionary events in the distant past. Incorporating them into phylogenetic hypotheses of living clades can help time-calibrate divergences, as well as elucidate macroevolutionary dynamics. However, the effect fossils have on phylogenetic reconstruction from morphology remains controversial. The consequences of explicitly incorporating the stratigraphic ages of fossils using tip-dated inference are also unclear. Here, we use simulations to evaluate the performance of inference methods across different levels of fossil sampling and missing data. Our results show that fossil taxa improve phylogenetic analysis of morphological datasets, even when highly fragmentary. Irrespective of inference method, fossils improve the accuracy of phylogenies and increase the number of resolved nodes. They also induce the collapse of ancient and highly uncertain relationships that tend to be incorrectly resolved when sampling only extant taxa. Furthermore, tip-dated analyses under the fossilized birth-death process outperform undated methods of inference, demonstrating that the stratigraphic ages of fossils contain vital phylogenetic information. Fossils help to extract true phylogenetic signals from morphology, an effect that is mediated by both their distinctive morphology and their temporal information, and their incorporation in total-evidence phylogenetics is necessary to faithfully reconstruct evolutionary history.

A Cambrian crown annelid reconciles phylogenomics and the fossil record.

The phylum of annelids is one of the most disparate animal phyla and encompasses ambush predators, suspension feeders and terrestrial earthworms1. The early evolution of annelids remains obscure or controversial2,3, partly owing to discordance between molecular phylogenies and fossils2,4. Annelid fossils from the Cambrian period have morphologies that indicate epibenthic lifestyles, whereas phylogenomics recovers sessile, infaunal and tubicolous taxa as an early diverging grade5. Magelonidae and Oweniidae (Palaeoannelida1) are the sister group of all other annelids but contrast with Cambrian taxa in both lifestyle and gross morphology2,6. Here we describe a new fossil polychaete (bristle worm) from the early Cambrian Canglangpu formation7 that we name Dannychaeta tucolus, which is preserved within delicate, dwelling tubes that were originally organic. The head has a well-defined spade-shaped prostomium with elongated ventrolateral palps. The body has a wide, stout thorax and elongated abdomen with biramous parapodia with parapodial lamellae. This character combination is shared with extant Magelonidae, and phylogenetic analyses recover Dannychaeta within Palaeoannelida. To our knowledge, Dannychaeta is the oldest polychaete that unambiguously belongs to crown annelids, providing a constraint on the tempo of annelid evolution and revealing unrecognized ecological and morphological diversity in ancient annelids.

Death is on Our Side: Paleontological Data Drastically Modify Phylogenetic Hypotheses.

Fossils are the only remaining evidence of the majority of species that have ever existed, providing a direct window into events in evolutionary history that shaped the diversification of life on Earth. Phylogenies underpin our ability to make sense of evolution but are routinely inferred using only data available from living organisms. Although extinct taxa have been shown to add crucial information for inferring macroevolutionary patterns and processes (such as ancestral states, paleobiogeography and diversification dynamics), the role fossils play in reconstructing phylogeny is controversial. Since the early years of phylogenetic systematics, different studies have dismissed the impact of fossils due to their incompleteness, championed their ability to overturn phylogenetic hypotheses or concluded that their behavior is indistinguishable from that of extant taxa. Based on taxon addition experiments on empirical data matrices, we show that the inclusion of paleontological data has a remarkable effect in phylogenetic inference. Incorporating fossils often (yet not always) induces stronger topological changes than increasing sampling of extant taxa. Fossils also produce unique topological rearrangements, allowing the exploration of regions of treespace that are never visited by analyses of only extant taxa. Previous studies have proposed a suite of explanations for the topological behavior of fossils, such as their retention of unique morphologies or their ability to break long branches. We develop predictive models that demonstrate that the possession of distinctive character state combinations is the primary predictor of the degree of induced topological change, and that the relative impact of taxa (fossil and extant) can be predicted to some extent before any phylogenetic analysis. Our results bolster the consensus of recent empirical studies by showing the unique role of paleontological data in phylogenetic inference, and provide the first quantitative assessment of its determinants, with broad consequences for the design of taxon sampling in both morphological and total-evidence analyses. [phylogeny, morphology, fossils, parsimony, Bayesian inference.].

Jaw elements in Plumulites bengtsoni confirm that machaeridians are extinct armoured scaleworms.

Machaeridians are Palaeozoic animals that are dorsally armoured with serialized, imbricating shell plates that cover or enclose the body. Prior to the discovery of an articulated plumulitid machaeridian from the Early Ordovician of Morocco that preserved unambiguous annelid characters (segmental parapodia with chaetae), machaeridians were a palaeontological mystery, having been previously linked to echinoderms, barnacles, tommotiids (putative stem-group brachiopods) or molluscs. Although the annelid affinities of machaeridians are now firmly established, their position within the phylum and relevance for understanding the early evolution of Annelida is less secure, with competing hypotheses placing Machaeridia in the stem or deeply nested within the crown group of annelids. We describe a scleritome of Plumulites bengtsoni from the Fezouata Formation of Morocco that preserves an anterior jaw apparatus consisting of at least two discrete elements that exhibit growth lines. Although jaws have multiple independent origins within the annelid crown group, comparable jaws are present only within Phyllodocida, the clade that contains modern aphroditiforms (scaleworms and relatives). Phylogenetic analysis places a monophyletic Machaeridia within the crown group of Phyllodocida in total-group Aphroditiformia, consistent with a common origin of machaeridian shell plates and scaleworm elytrae. The inclusion of machaeridians in Aphroditiformia truncates the ghost lineage of Phyllodocida by almost a hundred million years.

Cambrian Sessile, Suspension Feeding Stem-Group Ctenophores and Evolution of the Comb Jelly Body Plan.

The origin of ctenophores (comb jellies) is obscured by their controversial phylogenetic position, with recent phylogenomic analyses resolving either sponges or ctenophores as the sister group of all other animals. Fossil taxa can provide morphological evidence that may elucidate the origins of derived characters and shared ancestries among divergent taxa, providing a means to "break" long branches in phylogenetic trees. Here we describe new fossil material from the early Cambrian Chengjiang Biota, Yunnan Province, China, including the putative cnidarian Xianguangia, the new taxon Daihua sanqiong gen et sp. nov., and Dinomischus venustus, informally referred to as "dinomischids" here. "Dinomischids" possess a basal calyx encircled by 18 tentacles that surround the mouth. The tentacles carry pinnules, each with a row of stiff filamentous structures interpreted as very large compound cilia of a size otherwise only known in ctenophores. Together with the Cambrian tulip animal Siphusauctum and the armored Cambrian scleroctenophores, they exhibit anatomies that trace ctenophores to a sessile, polypoid stem lineage. This body plan resembles the polypoid, tentaculate morphology of cnidarians, including a blind gastric cavity partitioned by mesenteries. We propose that comb rows are derived from tentacles with paired sets of pinnules that each bear a row of compound cilia. The scleroctenophores exhibit paired comb rows, also observed in Siphusauctum, in addition to an organic skeleton, shared as well by Dinomischus, Daihua, and Xianguangia. We formulate a hypothesis in which ctenophores evolved from sessile, polypoid suspension feeders, sharing similarities with cnidarians that suggest either a close relationship between these two phyla, a striking pattern of early convergent evolution, or an ancestral condition for either metazoans or eumetazoans.

Bilateral Jaw Elements in Amiskwia sagittiformis Bridge the Morphological Gap between Gnathiferans and Chaetognaths.

Amiskwia sagittiformis Walcott 1911 is an iconic soft-bodied taxon from the Burgess Shale [1-3]. It was originally interpreted as a chaetognath [1], but it was later interpreted as a pelagic nemertean [2] or considered of uncertain affinity [3]. Part of this ambiguity is due to direct comparisons with members of the crown groups of extant phyla [4] and a lack of clarity regarding the systematic position of chaetognaths, which would allow for assessing character polarity in the phylum with respect to outgroups. Here, we show that Amiskwia preserves a bilaterally arranged set of head structures visible in relief and high reflectivity. These structures are best interpreted as jaws situated within an expanded pharyngeal complex. Morphological studies have highlighted a likely homology between bilateral and chitinous jaw elements in gnathiferans and chaetognaths [5], which is congruent with a shared unique Hox gene that suggests a close relationship between Gnathifera and Chaetognatha [6]. Molecular phylogenetic studies have recently found gnathiferans to be a deep branch of Spiralia and Chaetognaths either a sister group to Spiralia [7] or forming a clade with gnathiferans [6, 8]. Our phylogenetic analyses render Gnathifera paraphyletic with respect to Chaetognatha, and we therefore suggest that Amiskwia is best interpreted as a stem chaetognath, but crown gnathiferan.

Soft-Bodied Fossils Are Not Simply Rotten Carcasses - Toward a Holistic Understanding of Exceptional Fossil Preservation: Exceptional Fossil Preservation Is Complex and Involves the Interplay of Numerous Biological and Geological Processes.

Exceptionally preserved fossils are the product of complex interplays of biological and geological processes including burial, autolysis and microbial decay, authigenic mineralization, diagenesis, metamorphism, and finally weathering and exhumation. Determining which tissues are preserved and how biases affect their preservation pathways is important for interpreting fossils in phylogenetic, ecological, and evolutionary frameworks. Although laboratory decay experiments reveal important aspects of fossilization, applying the results directly to the interpretation of exceptionally preserved fossils may overlook the impact of other key processes that remove or preserve morphological information. Investigations of fossils preserving non-biomineralized tissues suggest that certain structures that are decay resistant (e.g., the notochord) are rarely preserved (even where carbonaceous components survive), and decay-prone structures (e.g., nervous systems) can fossilize, albeit rarely. As we review here, decay resistance is an imperfect indicator of fossilization potential, and a suite of biological and geological processes account for the features preserved in exceptional fossils.

Ichnological evidence for meiofaunal bilaterians from the terminal Ediacaran and earliest Cambrian of Brazil.

The evolutionary events during the Ediacaran-Cambrian transition (~541 Myr ago) are unparalleled in Earth history. The fossil record suggests that most extant animal phyla appeared in a geologically brief interval, with the oldest unequivocal bilaterian body fossils found in the Early Cambrian. Molecular clocks and biomarkers provide independent estimates for the timing of animal origins, and both suggest a cryptic Neoproterozoic history for Metazoa that extends considerably beyond the Cambrian fossil record. We report an assemblage of ichnofossils from Ediacaran-Cambrian siltstones in Brazil, alongside U-Pb radioisotopic dates that constrain the age of the oldest specimens to 555-542 Myr. X-ray microtomography reveals three-dimensionally preserved traces ranging from 50 to 600 µm in diameter, indicative of small-bodied, meiofaunal tracemakers. Burrow morphologies suggest they were created by a nematoid-like organism that used undulating locomotion to move through the sediment. This assemblage demonstrates animal-sediment interactions in the latest Ediacaran period, and provides the oldest known fossil evidence for meiofaunal bilaterians. Our discovery highlights meiofaunal ichnofossils as a hitherto unexplored window for tracking animal evolution in deep time, and reveals that both meiofaunal and macrofaunal bilaterians began to explore infaunal niches during the late Ediacaran.

Multiple optimality criteria support Ornithoscelida.

A recent study of early dinosaur evolution using equal-weights parsimony recovered a scheme of dinosaur interrelationships and classification that differed from historical consensus in a single, but significant, respect; Ornithischia and Saurischia were not recovered as monophyletic sister-taxa, but rather Ornithischia and Theropoda formed a novel clade named Ornithoscelida. However, these analyses only used maximum parsimony, and numerous recent simulation studies have questioned the accuracy of parsimony under equal weights. Here, we provide additional support for this alternative hypothesis using Bayesian implementation of the Mkv model, as well as through number of additional parsimony analyses, including implied weighting. Using Bayesian inference and implied weighting, we recover the same fundamental topology for Dinosauria as the original study, with a monophyletic Ornithoscelida, demonstrating that the main suite of methods used in morphological phylogenetics recover this novel hypothesis. This result was further scrutinized through the systematic exclusion of different character sets. Novel characters from the original study (those not taken or adapted from previous phylogenetic studies) were found to be more important for resolving the relationships within Dinosauromorpha than the relationships within Dinosauria. Reanalysis of a modified version of the character matrix that supports the Ornithischia-Saurischia dichotomy under maximum parsimony also supports this hypothesis under implied weighting, but not under the Mkv model, with both Theropoda and Sauropodomorpha becoming paraphyletic with respect to Ornithischia.

Earth's oldest 'Bobbit worm' - gigantism in a Devonian eunicidan polychaete.

Whilst the fossil record of polychaete worms extends to the early Cambrian, much data on this group derive from microfossils known as scolecodonts. These are sclerotized jaw elements, which generally range from 0.1-2 mm in size, and which, in contrast to the soft-body anatomy, have good preservation potential and a continuous fossil record. Here we describe a new eunicidan polychaete, Websteroprion armstrongi gen. et sp. nov., based primarily on monospecific bedding plane assemblages from the Lower-Middle Devonian Kwataboahegan Formation of Ontario, Canada. The specimens are preserved mainly as three-dimensional moulds in the calcareous host rock, with only parts of the original sclerotized jaw walls occasionally present. This new taxon has a unique morphology and is characterized by an unexpected combination of features seen in several different Palaeozoic polychaete families. Websteroprion armstrongi was a raptorial feeder and possessed the largest jaws recorded in polychaetes from the fossil record, with maxillae reaching over one centimetre in length. Total body length of the species is estimated to have reached over one metre, which is comparable to that of extant 'giant eunicid' species colloquially referred to as 'Bobbit worms'. This demonstrates that polychaete gigantism was already a phenomenon in the Palaeozoic, some 400 million years ago.

Uncertain-tree: discriminating among competing approaches to the phylogenetic analysis of phenotype data.

Morphological data provide the only means of classifying the majority of life's history, but the choice between competing phylogenetic methods for the analysis of morphology is unclear. Traditionally, parsimony methods have been favoured but recent studies have shown that these approaches are less accurate than the Bayesian implementation of the Mk model. Here we expand on these findings in several ways: we assess the impact of tree shape and maximum-likelihood estimation using the Mk model, as well as analysing data composed of both binary and multistate characters. We find that all methods struggle to correctly resolve deep clades within asymmetric trees, and when analysing small character matrices. The Bayesian Mk model is the most accurate method for estimating topology, but with lower resolution than other methods. Equal weights parsimony is more accurate than implied weights parsimony, and maximum-likelihood estimation using the Mk model is the least accurate method. We conclude that the Bayesian implementation of the Mk model should be the default method for phylogenetic estimation from phenotype datasets, and we explore the implications of our simulations in reanalysing several empirical morphological character matrices. A consequence of our finding is that high levels of resolution or the ability to classify species or groups with much confidence should not be expected when using small datasets. It is now necessary to depart from the traditional parsimony paradigms of constructing character matrices, towards datasets constructed explicitly for Bayesian methods.

The impact of fossil data on annelid phylogeny inferred from discrete morphological characters.

As a result of their plastic body plan, the relationships of the annelid worms and even the taxonomic makeup of the phylum have long been contentious. Morphological cladistic analyses have typically recovered a monophyletic Polychaeta, with the simple-bodied forms assigned to an early-diverging clade or grade. This is in stark contrast to molecular trees, in which polychaetes are paraphyletic and include clitellates, echiurans and sipunculans. Cambrian stem group annelid body fossils are complex-bodied polychaetes that possess well-developed parapodia and paired head appendages (palps), suggesting that the root of annelids is misplaced in morphological trees. We present a reinvestigation of the morphology of key fossil taxa and include them in a comprehensive phylogenetic analysis of annelids. Analyses using probabilistic methods and both equal- and implied-weights parsimony recover paraphyletic polychaetes and support the conclusion that echiurans and clitellates are derived polychaetes. Morphological trees including fossils depict two main clades of crown-group annelids that are similar, but not identical, to Errantia and Sedentaria, the fundamental groupings in transcriptomic analyses. Removing fossils yields trees that are often less resolved and/or root the tree in greater conflict with molecular topologies. While there are many topological similarities between the analyses herein and recent phylogenomic hypotheses, differences include the exclusion of Sipuncula from Annelida and the taxa forming the deepest crown-group divergences.

A new fireworm (Amphinomidae) from the Cretaceous of Lebanon identified from three-dimensionally preserved myoanatomy.

BACKGROUND: Rollinschaeta myoplena gen. et sp. nov is described from the Late Cretaceous (Cenomanian) Konservat-Lagerstätten of Hakel and Hjoula, Lebanon. The myoanatomy of the fossils is preserved in exceptional detail in three dimensions as calcium phosphate, allowing the musculature of the body wall, gut and parapodia to be reconstructed in detail. RESULTS: The major muscle groups of polychaetes can be identified in Rollinschaeta, including longitudinal muscle bands, circular muscles, oblique muscles, the parapodial muscle complex and the gut musculature, with a resolution sufficient to preserve individual fibres. To allow meaningful comparison with the phosphatized fossil specimens, extant polychaetes were stained with iodine and visualised using microCT. Rollinschaeta myoplena possesses two pairs of dorsal longitudinal muscles, dorsal and ventral circular muscles and a single pair of ventral longitudinal muscles. While six longitudinal muscle bands are known from other polychaete groups, their presence in combination with circular muscles is unique to Amphinomidae, allowing these fossils to be diagnosed to family level based solely on their myoanatomy. The elongate, rectilinear body and equally sized, laterally projecting parapodia of Rollinschaeta are found only within Amphinominae, demonstrating that the Cretaceous species is derived amongst Amphinomida. CONCLUSION: The uniquely preserved myoanatomy of Rollinschaeta has allowed diagnosis of a fossil annelid to subfamily level using microCT as a comparative tool for exploring myoanatomy in fossil and extant polychaetes. Our results demonstrate that fossilized muscles can provide systematically informative anatomical detail and that they should be studied when preserved.