Trilobite tridents demonstrate sexual combat at 400 Mya.

The Devonian trilobite Walliserops carries a remarkable anterior cephalic trident posing a challenge to functional interpretation. A unique teratological specimen of Walliserops trifurcatus showing four, rather than three tines, is inconsistent with possible hypotheses connecting the trident to feeding techniques and suggests a sexually selected function. Malformations in a variety of living organisms support this conclusion. Morphometric comparisons to similar structures used for intraspecific combat in dynastine beetles show that the trident occupies a comparable shape space consistent with the hypothesis that it was a sexual combat weapon, the oldest reported example of its kind. This lends further credibility to the idea that some trilobites may have been strongly sexually dimorphic.

The three-dimensionally articulated oral apparatus of a Devonian heterostracan sheds light on feeding in Palaeozoic jawless fishes.

Attempts to explain the origin and diversification of vertebrates have commonly invoked the evolution of feeding ecology, contrasting the passive suspension feeding of invertebrate chordates and larval lampreys with active predation in living jawed vertebrates. Of the extinct jawless vertebrates that phylogenetically intercalate these living groups, the feeding apparatus is well-preserved only in the early diverging stem-gnathostome heterostracans. However, its anatomy remains poorly understood. Here, we use X-ray microtomography to characterize the feeding apparatus of the pteraspid heterostracan Rhinopteraspis dunensis (Roemer, 1855). The apparatus is composed of 13 plates arranged approximately bilaterally, most of which articulate from the postoral plate. Our reconstruction shows that the oral plates were capable of rotating around the transverse axis, but likely with limited movement. It also suggests the nasohypophyseal organs opened internally, into the pharynx. The functional morphology of the apparatus in Rhinopteraspis precludes all proposed interpretations of feeding except for suspension/deposit feeding and we interpret the apparatus as having served primarily to moderate the oral gape. This is consistent with evidence that at least some early jawless gnathostomes were suspension feeders and runs contrary to macroecological scenarios that envisage early vertebrate evolution as characterized by a directional trend towards increasingly active food acquisition.

Survivorship dynamics of the flora of Devonian Angarida.

Devonian plants in Siberia present protracted pioneer succession. Research into the survivorship dynamics of early plant communities upon the palaeocontinent Angarida have demonstrated that transgression and volcanogenic nutrient influx were key to the survival of colonizing plants. Taxic proportions show that migrating taxa entered Angarida from the southwest, Kuznetsk and Minusinsk basins, dispersing across the continent in waves through central areas northwards. The patterns of dispersal are consistent throughout the Devonian. Increased nutrient load from the active pulses of the Viluy-Yakutsk Large Igneous Province, biogeomorphic ecosystem engineering and the increased biomass of Angaridan plants are assisted by Late Devonian transgression. These cumulative factors can be linked to the Late Devonian marine extinctions observed in Siberia.

Complex wound response mechanisms and phellogen evolution - insights from Early Devonian euphyllophytes.

We analyze the oldest fossil occurrences of wound-response periderm to characterize the development of wound responses in early tracheophytes. The origin of periderm production by a cambium (phellogen), an innovation with key roles in protection of inner plant tissues, is poorly explored; understanding periderm development in early tracheophytes can illuminate key aspects of this process. Anatomy of wound-response tissues is characterized in serial sections in a new Early Devonian (Emsian; c. 400 Ma) euphyllophyte from Quebec (Canada) - Nebuloxyla mikmaqiana sp. nov. - and compared to previously described euphyllophyte periderm from the same fossil locality to reconstruct periderm development. Characterizing development in these oldest periderm occurrences allows us to propose a model for the development of wound-response periderm in early tracheophytes: by phellogen activity that is poorly coordinated laterally but bifacial, producing secondary tissues initially outwardly and subsequently inwardly. The earliest occurrences of wound periderm pre-date the oldest known periderm produced systemically as a regular ontogenetic stage (canonical periderm), suggesting that periderm evolved initially as a wound-response mechanism. We hypothesize that canonical periderm evolved by exaptation of this wound sealing mechanism, whose deployment was triggered by tangential tensional stresses induced in the superficial tissues by vascular cambial growth from within.

An early snapshot of plant-herbivore interactions: Psilophyton diakanthon sp. nov. from the Early Devonian of Gaspé (Quebec, Canada).

PREMISE: Trimerophytes are a plexus of early tracheophytes that form the base of the euphyllophyte clade and, thus, represent the link between the earliest land plants and modern-day ferns, sphenophytes, and seed plants. As the best-characterized trimerophyte, the genus Psilophyton occupies a key position in the euphyllophyte fossil record. We describe a new Psilophyton species that has implications for the evolution of plant-animal interactions. METHODS: The fossil material is preserved by permineralization in the Lower Devonian (Emsian) Battery Point Formation (Québec, Canada) and was studied in serial sections using the cellulose acetate peel technique. RESULTS: Psilophyton diakanthon sp. nov. differs from other Psilophyton species in possessing fibers that form a discontinuous layer in the inner cortex and two distinct types of spinescent emergences whose anatomy and morphology are consistent with roles in anti-herbivore defense. CONCLUSIONS: Psilophyton diakanthon adds another species to an already diverse genus. Its two morphologically distinct types of spinescence suggest that herbivory was rampant in plant-animal interactions and demonstrate that anti-herbivory defenses had reached a previously unrecognized level of sophistication by 400 million years ago, in the Early Devonian.

Gilsonicaris from the Lower Devonian Hunsrück slate is a eunicidan annelid and not the oldest crown anostracan crustacean.

The Lower Devonian (Lower Emsian, -400 Myr) roof slates of the Hunsrück in southeastern Germany have delivered a highly diverse and exceptionally preserved marine fauna that provides a unique snapshot into the anatomy and ecology of a wide range of Palaeozoic animals. Several of the described taxa, however, remain enigmatic in their affinity, at least until new pyritized features hidden under the surface of the slate are revealed using X-ray radiography or micro-computed tomography (µCT). Here, we redescribe such an enigmatic fossil, the putative anostracan crustacean Gilsonicaris rhenanus Van Straelen, 1943. Using µCT scanning, we unveil unprecedented details of its anatomy, including a ventral oral opening and four pairs of recalcitrant jaw elements. These jaws are morphologically consistent with the scolecodonts of eunicidan polychaetes, which along with the gross anatomy of the body and head unambiguously identifies G. rhenanus as a polychaete rather than an arthropod. While this discovery firmly discards the Early Devonian record of crown anostracans in the fossil record, it adds a new record of eunicidan soft tissues, which are surprisingly rare considering the abundant microfossil record of scolecodonts.

Prototaxites reinterpreted as mega-rhizomorphs, facilitating nutrient transport in early terrestrial ecosystems.

The enigmatic fossil Prototaxites found in successions ranging from the Middle Ordovician to the Upper Devonian was originally described as having conifer affinity. The current debate, however, suggests that they probably represent gigantic algal-fungal symbioses. Our re-investigation of permineralized Prototaxites specimens from two localities, the Heider quarry in Germany and the Bordeaux quarry in Canada, reveals striking anatomical similarities with modern fungal rhizomorphs Armillaria mellea. We analysed extant fungal rhizomorphs and fossil Prototaxites through light microscopy of their anatomy, Fourier transform infrared spectroscopy, X-ray microscopy, and Raman spectroscopy. Based on these comparisons, we interpret the Prototaxites as fungi. The detailed preservation of cell walls and possible organelles seen in transverse sections of Prototaxites reveal that fossilization initiated while the organism was alive, inhibiting the collapse of delicate cellular structures. Prototaxites has been interpreted to grow vertically by many previous workers. Here we propose an alternative view that Prototaxites represents a complex hyphal aggregation (rhizomorph) that may have grown horizontally similar to modern complex aggregated mycelial growth forms, such as cords and rhizomorphs. Their main function was possibly to redistribute water and nutrition from nutrient-rich to nutrient-poor areas facilitating the expansion for early land plant communities.

Earliest record of transfer cells in Lower Devonian plants.

Key sources of information on the nature of early terrestrial ecosystems are the fossilized remains of plants and associated organic encrustations, which are interpreted as either biofilms, biological soil crusts or lichens. The hypothesis that some of these encrustations might be the remains of the thalloid gametophytes of embryophytes provided the stimulus for this investigation. Fossils preserved in charcoal were extracted from Devonian Period (Lochkovian Stage, c. 410-419 Myr old) sediments at a geological site in Shropshire (UK). Scanning electron micrographs (SEMs) of the fossils were compared with new and published SEMs of extant bryophytes and tracheophytes, respectively. One specimen was further prepared and imaged by transmission electron microscopy. Fossils of thalloid morphology were composed almost entirely of cells with labyrinthine ingrowths; these also were present in fossils of axial morphology where they were associated with putative food-conducting cells. Comparison with modern embryophytes demonstrates that these distinctive cells are transfer cells (TCs). Our fossils provide by far the earliest geological evidence of TCs. They also show that some organic encrustations are the remains of thalloid land plants and that these are possibly part of the life cycle of a newly recognized group of plants called the eophytes.

Piecing together the eophytes - a new group of ancient plants containing cryptospores.

The earliest evidence for land plants comes from dispersed cryptospores from the Ordovician, which dominated assemblages for 60 million years. Direct evidence of their parent plants comes from minute fossils in Welsh Borderland Upper Silurian to Lower Devonian rocks. We recognize a group that had forking, striated axes with rare stomata terminating in valvate sporangia containing permanent cryptospores, but their anatomy was unknown especially regarding conducting tissues. Charcoalified fossils extracted from the rock using HF were selected from macerates and observed using scanning electron microscopy. Promising examples were split for further examination and compared with electron micrographs of the anatomy of extant bryophytes. Fertile fossil axes possess central elongate cells with thick walls bearing globules, occasional strands and plasmodesmata-sized pores. The anatomy of these cells best matches desiccation-tolerant food-conducting cells (leptoids) of bryophytes. Together with thick-walled epidermal cells and extremely small size, these features suggest that these plants were poikilohydric. Our new data on conducting cells confirms a combination of characters that distinguish the permanent cryptospore-producers from bryophytes and tracheophytes. We therefore propose the erection of a new group, here named the Eophytidae (eophytes).

A protoxylem pathway to evolution of pith? An hypothesis based on the Early Devonian euphyllophyte Leptocentroxyla.

BACKGROUND AND AIMS: The Early Devonian (Emsian, 400-395 Ma) tracheophyte Leptocentroxyla tetrarcha Bickner et Tomescu emend. Tomescu et McQueen combines plesiomorphic Psilophyton-type tracheid thickenings with xylem architecture intermediate between the plesiomorphic basal euphyllophyte haplosteles and the complex actinosteles of Middle Devonian euphyllophytes. We document xylem development in Leptocentroxyla based on anatomy and explore its implications, which may provide a window into the evolution of pith. METHODS: Leptocentroxyla is preserved by permineralization in the Battery Point Formation (Quebec, Canada). Serial sections obtained using the cellulose acetate peel technique document branching pattern, anatomy of trace divergence to appendages, protoxylem architecture, and variations in tracheid size and wall thickening patterns. KEY RESULTS: Leptocentroxyla has opposite decussate pseudo-whorled branching and mesarch protoxylem, and represents the earliest instance of central histological differentiation in a euphyllophyte actinostele. Tracheids at the centre of xylem exhibit simplified Psilophyton-type wall thickenings and are similar in size (at the axis centre) or smaller than the surrounding metaxylem tracheids (at the centre of appendage traces). CONCLUSIONS: The position and developmental attributes of the simplified Psilophyton-type tracheids suggest they may have been generated by the protoxylem developmental pathway. This supports the delayed and shortened protoxylem differentiation hypothesis, which explains the evolution of pith by (1) delay in the onset of differentiation and lengthening of cell growth duration in a central protoxylem strand; and (2) shortening of the interval of differentiation of those tracheids, leading to progressive simplification (and eventual loss) of secondary wall thickenings, and replacement of tracheids with a central parenchymatous area. NAC domain transcription factors and their interactions with abscisic acid may have provided the regulatory substrate for the developmental changes that led to the evolution of pith. These could have been orchestrated by selective pressures associated with the expansion of early vascular plants into water-stresses upland environments.

Taxon sampling and alternative hypotheses of relationships in the euphyllophyte plexus that gave rise to seed plants: insights from an Early Devonian radiatopsid.

An abrupt transition in the fossil record separates Early Devonian euphyllophytes with a simple structure from a broad diversity of structurally complex Middle-Late Devonian plants. Morphological evolution and phylogeny across this transition are poorly understood due to incomplete sampling of the fossil record. We document a new Early Devonian radiatopsid and integrate it in analyses addressing euphyllophyte relationships. Anatomically preserved Emsian fossils (402-394 Ma) from the Battery Point Formation (Gaspé, Quebec, Canada) are studied in serial sections. The phylogenetic analysis is based on a matrix of 31 taxa and 50 characters emphasising vegetative morphology (41 discrete, nine continuous). The new plant, Kenrickia bivena gen. et sp. nov., is one of very few structurally complex euphyllophytes documented in the Early Devonian. Inclusion of Kenrickia overturns previously established phylogenetic relationships among Radiatopses, reiterating the need for increased density of Early Devonian taxon sampling. Kenrickia is recovered as the sister lineage to all other radiatopsids, a clade in which paraphyletic Stenokoleales led to a lignophyte clade where archaeopterids and seed plants fall into sister clades. Our results shed light on early euphyllophyte relationships and evolution, indicating early exploration of structural complexity by multiple lineages and reiterating the potential of a single origin of secondary growth in euphyllophytes.

A new phylogeny of the cladoxylopsid plexus: contribution of an early cladoxylopsid from the Lower Devonian (Emsian) of Quebec.

PREMISE: Cladoxylopsids formed Earth's earliest forests and gave rise to the ancestors of sphenopsids and ferns. Lower Devonian (Emsian) strata of the Battery Point Formation (Quebec, Canada) contain new anatomically preserved cladoxylopsids, one of which is described in this article. To assess the phylogenetic position of this fossil and address questions of cladoxylopsid phylogeny, we conducted a comprehensive phylogenetic study. METHODS: Permineralized axes were studied in serial sections using the cellulose acetate peel technique. We evaluated phylogenetic relationships among cladoxylopsids using a data set of 36 new morphological characters and 31 species, in parsimony-constrained analyses. RESULTS: We describe Adelocladoxis praecox gen. et sp. nov., a cladoxylopsid with small actinostelic axes bearing dichotomously branched, helically arranged ultimate appendages and fusiform sporangia. Adelocladoxis provides the oldest evidence of cladoxylopsid anatomy, including ultimate appendages and sporangia. In agreement with non-phylogenetic classification schemes, our phylogenetic analysis resolves a basal grade of iridopterids and a clade of cladoxylopsids s.s., which includes a pseudosporochnalean cladoxylopsid clade, a cladoxylalean cladoxylopsid clade, and Adelocladoxis. CONCLUSIONS: Our phylogenetic analysis illuminates aspects of tempo and mode of evolution in the cladoxylopsid plexus. Originating prior to the Emsian, cladoxylopsids reached global distribution by the Frasnian. Iridopterids and cladoxylopsids s.s. radiated in the Emsian-Eifelian. The sequence of character change recovered by our phylogeny supports a transition from actinostelic protosteles to dissected steles, associated with an increase in xylem rib number and medullation generating a central parenchymatous area.

The oldest Devonian circumpolar ray-finned fish?

Actinopterygians (ray-finned fishes) are the most diverse group of living fishes, but have a sparse Devonian fossil record restricted to low palaeolatitudes. Here we report a new actinopterygian from the Paraná Basin of Brazil, which occupied a circumpolar position in the Palaeozoic. Available geological evidence supports a Middle Devonian or older age for this taxon, which shares features of the mandibular symphysis with the latest Devonian Tegeolepis. A phylogenetic analysis resolves these two as sister taxa. This new record expands the palaeogeographic distribution of Devonian ray-fins and suggests that gaps in their fossil record might be filled by exploring poorly sampled high-latitude localities within the Malvinokaffric Realm.

Parametric transitions between bare and vegetated states in water-driven patterns.

Conditions for vegetation spreading and pattern formation are mathematically framed through an analysis encompassing three fundamental processes: flow stochasticity, vegetation dynamics, and sediment transport. Flow unsteadiness is included through Poisson stochastic processes whereby vegetation dynamics appears as a secondary instability, which is addressed by Floquet theory. Results show that the model captures the physical conditions heralding the transition between bare and vegetated fluvial states where the nonlinear formation and growth of finite alternate bars are accounted for by Center Manifold Projection. This paves the way to understand changes in biogeomorphological styles induced by man in the Anthropocene and of natural origin since the Paleozoic (Devonian plant hypothesis).

Archaeosporites rhyniensis gen. et sp. nov. (Glomeromycota, Archaeosporaceae) from the Lower Devonian Rhynie chert: a fungal lineage morphologically unchanged for more than 400 million years.

BACKGROUND AND AIMS: Structurally preserved arbuscular mycorrhizas from the Lower Devonian Rhynie chert represent core fossil evidence of the evolutionary history of mycorrhizal systems. Moreover, Rhynie chert fossils of glomeromycotan propagules suggest that this lineage of arbuscular fungi was morphologically diverse by the Early Devonian; however, only a small fraction of this diversity has been formally described and critically evaluated. METHODS: Thin sections, previously prepared by grinding wafers of chert from the Rhynie beds, were studied by transmitted light microscopy. Fossils corresponding to the description of Archaeospora spp. occurred in 29 slides, and were measured, photographed and compared with modern-day species in that genus. KEY RESULTS: Sessile propagules <85 µm in diameter, some still attached to a sporiferous saccule, were found in early land plant axes and the chert matrix; they developed, in a similar manner to extant Archaeospora, laterally or centrally within the saccule neck. Microscopic examination and comparison with extant fungi showed that, morphologically, the fossils share the characters used to circumscribe the genus Archaeospora (Glomeromycota; Archaeosporales; Archaeosporaceae). CONCLUSIONS: The fossils can be assigned with confidence to the extant family Archaeosporaceae, but because molecular analysis is necessary to place organisms in these taxa to present-day genera and species, they are placed in a newly proposed fossil taxon, Archaeosporites rhyniensis.

Morphological (and not anatomical or reproductive) features define early vascular plant phylogenetic relationships.

PREMISE: Perhaps the most rapid period of vascular plant evolution occurred during the Silurian-Devonian time interval. Yet, few quantitative analyses have established the extent to which anatomical, morphological, or reproductive features contributed to this episode of tracheophyte diversification. METHODS: Phylogenetic analyses were performed using a newly revised matrix of 54 characters (with 158 character states) of 37 of the best-preserved Paleozoic (predominantly Devonian) plants. Equisetum was included to determine whether it aligns with fossil sphenopsids or taxa collectively considered "ferns". The topology of the 54-character consensus tree was then compared to the topologies generated using only reproductive features (18 characters; 47 character states), only anatomical features (14 characters; 54 character states), only morphological features (22 characters; 57 character states), and the three pairwise combinations (e.g., anatomical and morphological characters). RESULTS: The new 54-character tree topology continued to identify a trimerophyte-euphyllophyte clade and a zosterophyllophyte-lycophyte clade emerging from a Cooksonia-rhyniophyte plexus. Equisetum aligned with fossil sphenopsids rather than fern-like fossil taxa. Reproductive characters or anatomical characters analyzed in isolation resulted in nearly complete polytomy. Among the various permutations of the three categories, anatomical and morphological characters when combined provided the best restoration of the 54-character tree topology. CONCLUSIONS: The phylogenetic relationships among the canonical fossil taxa used in this analysis predominantly reflect morphological trends. Reproductive and anatomical features taken in isolation appear to be evolutionarily conservative characters, i.e., natural selection "sees" the external phenotype.

Unique growth strategy in the Earth's first trees revealed in silicified fossil trunks from China.

Cladoxylopsida included the earliest large trees that formed critical components of globally transformative pioneering forest ecosystems in the Mid- and early Late Devonian (ca. 393-372 Ma). Well-known cladoxylopsid fossils include the up to ∼1-m-diameter sandstone casts known as Eospermatopteris from Middle Devonian strata of New York State. Cladoxylopsid trunk structure comprised a more-or-less distinct cylinder of numerous separate cauline xylem strands connected internally with a network of medullary xylem strands and, near the base, externally with downward-growing roots, all embedded within parenchyma. However, the means by which this complex vascular system was able to grow to a large diameter is unknown. We demonstrate-based on exceptional, up to ∼70-cm-diameter silicified fossil trunks with extensive preservation of cellular anatomy from the early Late Devonian (Frasnian, ca. 374 Ma) of Xinjiang, China-that trunk expansion is associated with a cylindrical zone of diffuse secondary growth within ground and cortical parenchyma and with production of a large amount of wood containing both rays and growth increments concentrically around individual xylem strands by normal cambia. The xylem system accommodates expansion by tearing of individual strand interconnections during secondary development. This mode of growth seems indeterminate, capable of producing trees of large size and, despite some unique features, invites comparison with secondary development in some living monocots. Understanding the structure and growth of cladoxylopsids informs analysis of canopy competition within early forests with the potential to drive global processes.

The early elasmobranch Phoebodus: phylogenetic relationships, ecomorphology and a new time-scale for shark evolution.

Anatomical knowledge of early chondrichthyans and estimates of their phylogeny are improving, but many taxa are still known only from microremains. The nearly cosmopolitan and regionally abundant Devonian genus Phoebodus has long been known solely from isolated teeth and fin spines. Here, we report the first skeletal remains of Phoebodus from the Famennian (Late Devonian) of the Maïder region of Morocco, revealing an anguilliform body, specialized braincase, hyoid arch, elongate jaws and rostrum, complementing its characteristic dentition and ctenacanth fin spines preceding both dorsal fins. Several of these features corroborate a likely close relationship with the Carboniferous species Thrinacodus gracia, and phylogenetic analysis places both taxa securely as members of the elasmobranch stem lineage. Identified as such, phoebodont teeth provide a plausible marker for range extension of the elasmobranchs into the Middle Devonian, thus providing a new minimum date for the origin of the chondrichthyan crown-group. Among pre-Carboniferous jawed vertebrates, the anguilliform body shape of Phoebodus is unprecedented, and its specialized anatomy is, in several respects, most easily compared with the modern frilled shark Chlamydoselachus. These results add greatly to the morphological, and by implication ecological, disparity of the earliest elasmobranchs.

Bite marks and predation of fossil jawless fish during the rise of jawed vertebrates.

Although modern vertebrate diversity is dominated by jawed vertebrates, early vertebrate assemblages were predominantly composed of jawless fishes. Hypotheses for this faunal shift and the Devonian decline of jawless vertebrates include predation and competitive replacement. The nature and prevalence of ecological interactions between jawed and jawless vertebrates are highly relevant to both hypotheses, but direct evidence is limited. Here, we use the occurrence and distribution of bite mark type traces in fossil jawless armoured heterostracans to infer predation interactions. A total of 41 predated specimens are recorded; their prevalence increases through time, reaching a maximum towards the end of the Devonian. The bite mark type traces significantly co-occur with jawed vertebrates, and their distribution through time is correlated with jawed vertebrate diversity patterns, particularly placoderms and sarcopterygians. Environmental and ecological turnover in the Devonian, especially relating to the nekton revolution, have been inferred as causes of the faunal shift from jawless to jawed vertebrates. Here, we provide direct evidence of escalating predation from jawed vertebrates as a potential contributing factor to the demise and extinction of ostracoderms.

Belowground rhizomes in paleosols: The hidden half of an Early Devonian vascular plant.

The colonization of terrestrial environments by rooted vascular plants had far-reaching impacts on the Earth system. However, the belowground structures of early vascular plants are rarely documented, and thus the plant-soil interactions in early terrestrial ecosystems are poorly understood. Here we report the earliest rooted paleosols (fossil soils) in Asia from Early Devonian deposits of Yunnan, China. Plant traces are extensive within the soil and occur as complex network-like structures, which are interpreted as representing long-lived, belowground rhizomes of the basal lycopsid Drepanophycus The rhizomes produced large clones and helped the plant survive frequent sediment burial in well-drained soils within a seasonal wet-dry climate zone. Rhizome networks contributed to the accumulation and pedogenesis of floodplain sediments and increased the soil stabilizing effects of early plants. Predating the appearance of trees with deep roots in the Middle Devonian, plant rhizomes have long functioned in the belowground soil ecosystem. This study presents strong, direct evidence for plant-soil interactions at an early stage of vascular plant radiation. Soil stabilization by complex rhizome systems was apparently widespread, and contributed to landscape modification at an earlier time than had been appreciated.