Different analytical methods can yield competing interpretations of evolutionary history and, currently, there is no definitive method for phylogenetic reconstruction using morphological data. Parsimony has been the primary method for analysing morphological data, but there has been a resurgence of interest in the likelihood-based Mk-model. Here, we test the performance of the Bayesian implementation of the Mk-model relative to both equal and implied-weight implementations of parsimony. Using simulated morphological data, we demonstrate that the Mk-model outperforms equal-weights parsimony in terms of topological accuracy, and implied-weights performs the most poorly. However, the Mk-model produces phylogenies that have less resolution than parsimony methods. This difference in the accuracy and precision of parsimony and Bayesian approaches to topology estimation needs to be considered when selecting a method for phylogeny reconstruction.

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.

Highlights d We describe a new early Cambrian fossil with a polypoid body plan from Chengjiang d It is related to the supposed cnidarian Xianguangia and the iconic Dinomischus d The fossils possess very large ciliary structures otherwise seen only in ctenophores d We show that these fossils form a grade of stem-group ctenophores

Exceptionally preserved fossils provide crucial insights into extinct body plans and organismal evolution(1). Molluscs, one of the most disparate animal phyla, radiated rapidly during the early Cambrian period (approximately 535-520 million years ago (Ma))(2). The problematic fossil taxa Halkieria(3) and Orthrozanclus(4) (grouped in Sachitida) have been assigned variously to stem-group annelids, brachiopods(4,5), stem-group molluscs(4) or stem-group aculiferans (Polyplacophora and Aplacophora)(6), but their affinities have remained controversial owing to a lack of preserved diagnostic characters. Here we describe a new early sachitid, Calvapilosa kroegeri gen. et sp. nov. from the Fezouata biota of Morocco(7,8) (Early Ordovician epoch, around 478 Ma). The new taxon is characterized by the presence of a single large anterior shell plate and polystichous radula bearing a median tooth and several lateral and uncinal teeth in more than 125 rows. Its flattened body is covered by hollow spinose sclerites, and a smooth, ventral girdle flanks an extensive mantle cavity. Phylogenetic analyses resolve C. kroegeri as a stem-group aculiferan together with other single-plated forms such as Maikhanella (Siphogonuchites) and Orthrozanclus; Halkieria is recovered closer to the aculiferan crown. These genera document the stepwise evolution of the aculiferan body plan from forms with a single, almost conchiferan-like shell through two-plated taxa such as Halkieria, to the eight-plated crown-group aculiferans. C. kroegeri therefore provides key evidence concerning the long debate about the crown molluscan affinities of sachitids. This new discovery strongly suggests that the possession of only a single calcareous shell plate and the presence of unmineralised sclerites are plesiomorphic (an ancestral trait) for the molluscan crown

Annelids are a phylum of segmented bilaterian animals that have become important components of ecosystems spanning terrestrial realms to the deep sea. Annelids are remarkably diverse, possessing high taxonomic diversity and exceptional morphological disparity, and have evolved numerous feeding strategies and ecologies. Their interrelationships and evolution have been the source of much controversy over the past century with the composition of the annelid crown group, the relationship of major groups and the body plan of the ancestral annelid having undergone major recent revisions. There is a convincing body of molecular evidence that polychaetes form a paraphyletic grade and that clitellates are derived polychaetes. The earliest stem group annelids from Cambrian Lagerst€ atten are errant, epibenthic polychaetes, confirming that biramous parapodia, head appendages and diverse, simple chaetae are primitive for annelids. Current evidence from molecular clocks and the fossil record suggest that crown group annelids are a Late Cambrian -Ordovician radiation, with clitellates radiating in the Late Palaeozoic. Their body fossil record is largely confined to deposits showing exceptional preservation and is punctuated by the acquisition of hard parts in major groups. The discovery of an Ordovician fossil with soft tissues has shown that machaeridians are in fact a clade of crown polychaetes. They were in existence for more than 200 million years and possess unique calcitic dorsal armour, allowing their mode of life and phylogeny to be interpreted in the context of the annelid body plan. We identify a novel clade of machaeridians, the Cuniculepadida, which exhibit a series of adaptations for burrowing.

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 μ m 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. NATuRE ECoLoGy & EvoLuTIoN Articles Nature ecology & evolutioNsupport for these suggestions is limited to purported body fossils of sponges 19 and demosponge biomarkers 20 . A considerable gap therefore remains between the fossil record of the late Ediacaran and molecular clock estimates for deep splits in the animal tree, for example the origin of Metazoa and Eumetazoa 3 . Assuming that contemporary molecular clock analyses yield accurate, if imprecise 18 , node ages for animal divergences, a small body size and concomitant limited fossilization potential 21 could reconcile these discordant records of animal evolution (but see ref. 22 ).The small body size of the ancestral bilaterian is supported by recent phylogenomic analyses of deep animal relationships, with acoel flatworms and xenoturbellids (Xenacoelomorpha) being a sister group to all remaining bilaterians (Nephrozoa) 23 , and smallbodied spiralian taxa (the 'Platyzoa') recognized as a paraphyletic grade with respect to macroscopic trochozoans 24 . This suggests that early bilaterians and spiralians were small bodied, possibly meiofaunal, and moved using ciliary gliding.Meiofauna comprises all organisms between 32 and 1,000 μ m in size that inhabit pore-water-rich sediments in freshwater to deepmarine environments 25 . Modern meiofaunal communities include animals, foraminifera and some ciliates, and contribute significantly to sediment bioturbation and bioirrigation 26,27 . The meiofauna can be divided into permanent members (that is, animals with organisms of a small size adapted and restricted to the meiofaunal, interstitial realm) and temporary meiofauna (for example, the ...

A significant negative relationship between tiller height and resistance to Fusarium ear blight (FEB), following inoculation, was observed in 17 cultivars of winter wheat in the 1995/96 growing season. Tall cultivars such as Kraka and Spark showed fewer symptoms of FEB (6 and 4%) than the shorter cultivars Brigadier and Virtue (35 and 51%). To determine if this relationship was caused by a genetic association or an effect of the microclimate, height and disease were measured in segregating populations derived from tall × short cultivars and humidity was measured in near-isogenic lines with and without the Rht1 and Rht2 dwarfing genes. Among random F 3 populations there was a clear tendency for tall strawed lines to show less severe disease symptoms than shorter strawed lines following inoculation. The effect of the individual dwarfing genes Rht1 and Rht2 on the severity of FEB was also studied in an inoculation trial using a number of near-isogenic lines of Maris Huntsman and Maris Widgeon. Within isogenic lines of Maris Huntsman, there was a clear tendency for tall straw to be associated with fewer symptoms, but this was not apparent within lines of Maris Widgeon. Monitoring relative humidity at ear height in a short and tall isogenic line of Maris Huntsman revealed no significant differences between these genotypes from GS 65 to GS 85, suggesting that microclimate cannot explain differences in severity of FEB between these lines. It is suggested that there are independent genes affecting the severity of FEB that may allow plant breeders to select resistant cultivars of any height.

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.