Zirabia cylindrica comb. nov. provides evidence of Doyleales in the Jurassic.

PREMISE: In recent years, Doyleales have played an important role in the discussion and exploration of seed plant relationships and the origin of angiosperms. This order comprises a diversity of recently described genera with cupule-bearing compound ovulate cones recovered from Early Cretaceous deposits in North America and Asia. Their relatively late appearance in the fossil record, stratigraphically near the appearance of angiosperms in the Early Cretaceous, has been noteworthy. Here, we report a new genus of Doyleales, Zirabia gen. nov. from the Early Jurassic of Iran, that was originally described as the ginkgophyte Karkenia. METHODS: We reinvestigated material previously assigned to Karkenia cylindrica from the Lower Jurassic of the Zirab locality, Alborz Mountains, northern Iran. RESULTS: The studied specimen features a main axis bearing helically to irregularly arranged bract-cupule complexes, each composed by a long laminar bract subtending and sheathing a cupule stalk that bears a single-seeded cupule with a dorsal protrusion. The morphological features of this taxon do not conform with those of Karkenia, and suggest affinities with Doyleales rather than Ginkgoales. Within Doyleales, this fossil has a unique combination of characters indicating that it is a new genus within the order; thus, a new combination is erected, Zirabia cylindrica. CONCLUSIONS: Our results indicate that Doyleales is significantly older than previously thought, with their stratigraphic range now extending from the Lower Jurassic to the Cretaceous. The Early Jurassic occurrence of Doyleales provides important data on the emergence of seed-enclosing structures seen in seed plants throughout the Mesozoic.

A phylogenomic analysis of Lonicera and its bearing on the evolution of organ fusion.

PREMISE: The ~140 species of Lonicera are characterized by variously fused leaves, bracteoles, and ovaries, making it a model system for studying the evolution and development of organ fusion. However, previous phylogenetic analyses, based mainly on chloroplast DNA markers, have yielded uncertain and conflicting results. A well-supported phylogeny of Lonicera will allow us to trace the evolutionary history of organ fusion. METHODS: We inferred the phylogeny of Lonicera using restriction site-associated DNA sequencing (RADSeq), sampling all major clades and 18 of the 23 subsections. This provided the basis for inferring the evolution of five fusion-related traits. RESULTS: RADSeq data yielded a well-resolved and well-supported phylogeny. The two traditionally recognized subgenera (Periclymenum and Chamaecerasus), three of the four sections (Isoxylosteum, Coeloxylosteum, and Nintooa), and half of the subsections sampled were recovered as monophyletic. However, the large and heterogeneous section Isika was strongly supported as paraphyletic. Nintooa, a clade of ~22 mostly vine-forming species, including L. japonica, was recovered in a novel position, raising the possibility of cytonuclear discordance. We document the parallel evolution of fused leaves, bracteoles, and ovaries, with rare reversals. Most strikingly, complete cupules, in which four fused bracteoles completely enclose two unfused ovaries, arose at least three times. Surprisingly, these appear to have evolved directly from ancestors with free bracteoles instead of partial cupules. CONCLUSIONS: We provide the most comprehensive and well-supported phylogeny of Lonicera to date. Our inference of multiple evolutionary shifts in organ fusion provides a solid foundation for in depth developmental and functional analyses.

A novel cupulate seed plant, Xadzigacalix quatsinoensis gen. et sp. nov., provides new insight into the Mesozoic radiation of gymnosperms.

PREMISE: Anatomically preserved evidence for a novel clade of gymnosperms emphasizes diversity of seed plants immediately prior to the appearance of angiosperm fossils in the paleontological record. METHODS: Cupulate seeds from the Early Cretaceous Apple Bay locality (Vancouver Island) are described from serial cellulose acetate peels and three-dimensional reconstruction. Phylogenetic context is assessed through the comparative analysis of gymnosperm seed producing fructifications and maximum parsimony analysis of a revised morphological data set for seed plant phylogeny. RESULTS: Xadzigacalix quatsinoensis gen. et sp. nov. is characterized by an orthotropous ovule with an elongated micropyle and complex integument, enclosed within a radial cupule. The micropylar canal is elongated; and the nucellus extends into the micropyle to seal the post pollination ovule. Except at the apex of the micropyle, the seed is completely enclosed by a parenchymatous cupule with ca. 20 axially elongated secretory ducts. The cupulate seed is produced upon a triangular woody stele, consisting of a parenchymatous pith surrounded by radially aligned tracheids. The stele produces three short terete traces that terminate within the base of the cupule as transfusion tissue at the seed chalaza. CONCLUSIONS: Organography, vascularization, nature of the integument and nucellus, and configuration of the micropylar canal distinguish Xadzigacalix quatsinoensis from all other gymnosperm clades. Cladistic analyses suggest the new plant may have affinities with gnetophytes or angiosperms. These results are complemented with a critical re-evaluation of ovulate structures for Mesozoic gymnosperms, providing new insight into plant diversity immediately antecedent to the explosive diversification of flowering plants.

Reconstructing development of the earliest seed integuments raises a new hypothesis for the evolution of ancestral seed-bearing structures.

How plant seeds originated remains unresolved, in part due to disconnects between fossil intermediates and developmental genetics in extant species. The Carboniferous fossil Genomosperma is considered among the most primitive known seeds, with highly lobed integument and exposed nucellus. We have used this key fossil taxon to investigate the evolutionary origins of seed development. We examined sectioned Genomosperma specimens using modern digital 3D reconstruction techniques and established population-level measurements of Genomosperma ovules for quantitative analysis. Genomosperma ovules show significant variation in integumentary lobe fusion and curvature. Our analysis suggests that this variation represents a single species with significant variations in lobe number and fusion, reminiscent of floral development in extant species. We conclude that changes in lobe flexure occurred late in development, consistent with a previously hypothesized function in pollen guidance/retention. We also identify seeds of Genomosperma within cupules for the first time. The presence of a cupule adds evidence towards the plesiomorphy of cupules within seed plants. Together with the similarities identified between the Genomosperma lobed integument and floral organs, we propose that the cupule, integument and nucellus together developed in a shoot-like fashion, potentially ancestral to extant seed plant reproductive shoots.

Fruit development of Lithocarpus (Fagaceae) and the role of heterochrony in their evolution.

Stone oaks, or Lithocarpus species of Fagaceae are ecologically important canopy trees in the tropical and subtropical forests over East Asia, and the fruits of which are important food sources for insects and vertebrates there. The great fruit morphological variation of this genus represents two fruit types, acorn and enclosed receptacle fruit types. However, the evolutionary mechanisms of differentiation into these two fruit types with contrasting morphology remain a puzzle. To reveal the morphogenetic properties of two fruit types, we observed tissue differentiation and development among 20 Lithocarpus species from fruit set to maturity. Unlike in fruits of Quercus, the endocarp differentiation in Lithocarpus fruits occurred later than exocarp and mesocarp. Cupules provided further protection of developing seeds, particularly of acorn-type fruits. Fruits of Lithocarpus and Quercus acorns share similar insect predators. At fruit set, both acorn and enclosed receptacle types were largely identical, with similar tissue morphology and the sequence of differentiation. The distinct difference between two fruit types at maturity came from varied rates and degrees of development between the pericarp and receptacle tissues. We found that heterochrony between two tissues could create substantially divergent ecological strategies for protection and dispersal of their seeds, which is essential for the evolution of two fruit types.

A late Cretaceous fagalean inflorescence preserved in amber from New Jersey.

PREMISE OF THE STUDY: An inflorescence with three pistillate flowers in amber from the early Upper Cretaceous (Turonian, ~90-94 million years ago) of central New Jersey represents the oldest known flowers with features present in an early stem complex of the Fagales. The inflorescence has characteristics of Nothofagaceae, but also has strikingly distinct characters that suggest it is intermediate between Nothofagus and other Fagales. This intermediacy is consistent with its northern hemisphere distribution. METHODS: We investigated this new fossil by comparing it with extant and fossil members of the Fagales using light microscopy and nano-computed tomography. In addition, for exploring its relationships, we mapped the morphological characters onto a widely accepted molecular-based tree of modern basal Fagales using standard methods of character optimization. KEY RESULTS: The phylogenetic position of the fossil inflorescence can be unequivocally determined by the presence of unique features, singly and in combination, that are found only in "basal" members of Fagales. The fossil adds critical information on the features of the early stem Fagales, evolution of the cupule in Nothofagaceae and Fagaceae, and a reasonable biogeographic hypothesis for the differentiation of southern (e.g., Nothofagaceae) and northern hemisphere Fagales. CONCLUSIONS: This new fossil provides insight into the early evolution of Fagales and suggests that early stem Fagales that had not yet differentiated into modern families were present in the Late Cretaceous of North America. Based on available evidence, the fossil is best interpreted as an early stem member of the Fagales, with features that suggest a transition from a more generalized Nothofagus-like fagalean ancestor with some unique presumably plesiomorphic features. The presence of an enlarged perianth and flexuous styles also suggests the possibility of insect pollination, which has been lost in all Fagales with the exception of some members of subfamily Castaneoideae in Fagaceae sensu stricto (which otherwise are very different from this fossil). The poorly developed, bract-like cupule valves of the fossil can be interpreted as primitive (i.e., incipient) or as reduced from more developed cupules that are found in most modern Fagaceae and Nothofagaceae.

Synchrotron X-ray imaging of a dichasium cupule of Castanopsis from Eocene Baltic amber.

PREMISE OF THE STUDY: The Eocene Baltic amber deposit represents the largest accumulation of fossil resin worldwide, and hundreds of thousands of entrapped arthropods have been recovered. Although Baltic amber preserves delicate plant structures in high fidelity, angiosperms of the "Baltic amber forest" remain poorly studied. We describe a pistillate partial inflorescence of Castanopsis (Fagaceae), expanding the knowledge of Fagaceae diversity from Baltic amber. METHODS: The amber specimen was investigated using light microscopy and synchrotron-radiation-based X-ray micro-computed tomography (SRµCT). KEY RESULTS: The partial inflorescence is a cymule, consisting of an involucre of scales that surround all four pistillate flowers, indicating a dichasium cupule. Subtending bracts are basally covered with peltate trichomes. Flowers possess an urecolate perianth of six nearly free lobes, 12 staminodia hidden by the perianth, and a tri-locular ovary that is convex-triangular in cross section. The exceptional three-dimensional preservation suggests that the fossil belongs to the extant East Asian genus Castanopsis. The amber inclusion represents the first record of Castanopsis from Baltic amber and the first pistillate inflorescence of Fagaceae from Eurasia. CONCLUSIONS: The partial female inflorescence reported here provides an important addition to acorns of Castanopsis described from middle Eocene strata of Europe. Furthermore, the intercontinental distribution of Castanopsis in the Eocene is confirmed. The amber fossil also broadens the picture of the Baltic amber source area, indicating oligotrophic, sandy, bog-like habitats. Finally, this study underscores the great benefit of SRµCT as a powerful tool to investigate plant inclusions from amber in a nondestructive way.

Latisemenia longshania, gen. et sp. nov., a new Late Devonian seed plant from China.

The earliest known ovules in the Late Devonian (Famennian) are borne terminally on fertile branches and are typically enclosed in a cupule. Among these ovules are some that have terete integumentary lobes with little or no fusion. Here, we report a new taxon, Latisemenia longshania, from the Famennian of South China, which bears cupulate ovules that are terminal as well as opposite on the fertile axis. Each ovule has four broad integumentary lobes, which are extensively fused to each other and also to the nucellus. The cupule is uniovulate, and the five flattened cupule segments of each terminal ovule are elongate cuneate and shorter than the ovule. Associated but not attached pinnules are laminate and Sphenopteris-like, with an entire or lobate margin. Latisemenia is the earliest known plant with ovules borne on the side of the fertile axis and may foreshadow the diverse ovule arrangements found among younger seed plant lineages that emerge in the Carboniferous. Following the telome theory, Latisemenia demonstrates derived features in both ovules and cupules, and the shape and fusion of integumentary lobes suggest effective pollination and protection to the nucellus. Along with other recent discoveries from China, Latisemenia extends the palaeogeographic range of the earliest seed plants.

Guazia, the earliest ovule without cupule but with unique integumentary lobes.

The earliest ovules in the Late Devonian (Famennian) are surrounded by a cupule that is involved in both protection and pollination, and generally have free integumentary lobes. Here we report a new taxon from the Famennian of China, Guazia dongzhiensis gen. et sp. nov. The terminally borne ovule is apparently acupulate (without cupule) and has four radially arranged wing-like integumentary lobes that are extensively fused, and folded lengthwise and inwards. Guazia provides evidence that not all Devonian seeds possess a cupule and it increases their diversity in integumentary lobes. This genus also suggests that the integuments develop new functions, probably including wind dispersal at the expense of the cupules.

Assessment of trapezial prosthetic cup migration: A biomechanical study.

We performed a biomechanical study using 60 Sawbones® rigid foam blocks of two simulated densities (osteoporotic, n = 30 and non-osteoporotic, n = 30) and 10 cadaveric trapezium bones from fresh-frozen, unembalmed adult cadaver hands to assess the trapezial prosthetic cup migration with progressively greater compression loads (10-40 kg). Two cups from the Touch® prosthesis were compared: 9-mm conical cup and 9-mm spherical cup. Uniaxial compression tests were carried out using an MTS Criterion® Series 40 Electromechanical Testing System. Cup migration was measured in millimeters (mm) at 10, 20, and 40 kg of compression load. Median cup migration values were similar in the cadaveric trapezium bones and Sawbones® non-osteoporotic blocks, and higher in the Sawbones® osteoporotic blocks. In the cadaveric trapezium bones and the Sawbones® non-osteoporotic blocks, migration values were less than or equal to 0.1 mm for 10 and 20 kg loads; it was 0.2 mm for 40 kg load. In the Sawbones® osteoporotic blocks, migration values were less than or equal to 0.3 mm for 10 and 20 kg loads; it was 0.4-0.5 mm for 40 kg load. There was no significant difference between the two cup shapes in both cadaveric trapezium bones and Sawbones® non-osteoporotic blocks. In Sawbones® osteoporotic blocks, the largest difference between the two cup shapes was 0.1 mm for loads up to 40 kg, which corresponded to our measurement accuracy. Our findings indicate that the trapezial component of total trapeziometacarpal joint arthroplasty undergoes very weak migration for axial compression loads up to 40 kg, presumably below the threshold of clinical relevance. The cup shape did not have an obvious influence; however, low bone mineral density may result in greater cup migration.

Comparison of two radiographic landmarks for centering the trapezial component in total trapeziometacarpal arthroplasty.

We performed a retrospective review of standard anteroposterior and lateral radiographs of the thumb in 80 patients, to compare two radiographic landmarks, in terms of mediolateral bone support, for centering the trapezial component in total joint arthroplasty. On anteroposterior view, we identified the distal articular surface of the trapezium and trapezium width, and defined the two midpoints as radiographic landmarks for positioning a 9-mm trapezial cup. Mean trapezium width was significantly greater than the distal articular surface of the trapezium, and the midpoints did not match. Thus, after positioning simulated 9-mm prosthetic cups centered on each landmark, the residual radial bone distance was significantly greater using the landmark based on trapezium width. The mean value was 33% greater with this landmark, and the minimum value was 2.1 mm, compared to 0.2 mm using the landmark based on the distal articular surface. Our study thus suggested that the midpoint of the trapezium width is the more relevant radiographic landmark for centering the trapezial prosthetic cup in total joint arthroplasty, by preserving better bone stock on the radial side without depleting the ulnar side. On an intraoperative anteroposterior fluoroscopic view, this landmark could be used to check cup positioning.

Development and evolution of the unique ovules of flowering plants.

Ovules are the precursors to seeds and as such are critical to plant propagation and food production. Mutant studies have led to the identification of numerous genes regulating ovule development. Genes encoding transcription factors have been shown to direct ovule spacing, ovule identity and integument formation. Particular co-regulators have now been associated with activities of some of these transcription factors, and other protein families including cell surface receptors have been shown to regulate ovule development. Hormone levels and transport, especially of auxin, have also been shown to play critical roles in ovule emergence and morphogenesis and to interact with the transcriptional regulators. Ovule diversification has been studied using orthologs of regulatory genes in divergent angiosperm groups. Combining modern genetic evidence with expanding knowledge of the fossil record illuminates the possible origin of the unique bitegmic ovules of angiosperms.

Beech cupules as keystone structures for soil fauna.

Facilitative or positive interactions are ubiquitous in nature and play a fundamental role in the configuration of ecological communities. In particular, habitat modification and niche construction, in which one organism locally modifies abiotic conditions and favours other organisms by buffering the effects of adverse environmental factors, are among the most relevant facilitative interactions. In line with this, 'keystone structures', which provide resources, refuge, or advantageous services decisive for other species, may allow the coexistence of various species and thus considerably contribute to diversity maintenance. Beech cupules are woody husks harbouring beech fruits that remain in the forest soil for relatively long periods of time. In this study, we explored the potential role of these cupules in the distribution and maintenance of the soil fauna inhabiting the leaf litter layer. We experimentally manipulated cupule availability and soil moisture in the field to determine if such structures are limiting and can provide moist shelter to soil animals during drought periods, contributing to minimize desiccation risks. We measured invertebrate abundances inside relative to outside the cupules, total abundances in the leaf litter and animal body sizes, in both dry and wet experimental plots. We found that these structures are preferentially used by the most abundant groups of smaller soil animals-springtails, mites and enchytraeids-during droughts. Moreover, beech cupules can be limiting, as an increase in use was found with higher cupule densities, and are important resources for many small soil invertebrates, driving the spatial structure of the soil community and promoting higher densities in the leaf litter, probably through an increase in habitat heterogeneity. We propose that fruit woody structures should be considered 'keystone structures' that contribute to soil community maintenance. Therefore, beech trees may indirectly facilitate soil fauna activities through their decaying fruit husks, hence acting as ecosystem engineers.

Retrospective study of two fixation methods for 4-corner fusion: Shape-memory staple vs. dorsal circular plate.

The purpose of this study was to compare the results of two groups of patients with four-corner fusion, one group fixed with shape-memory staples and the other with locked circular plates. This retrospective study compared 52 wrists operated for scaphoid excision and four-corner fusion between 2005 and 2011. The arthrodesis was ensured by a shape-memory quadripodal staple (4Fusion(®), Memometal™) in 37 cases and a locking dorsal circular plate (Xpode(®), Biotech Ortho™) in 15 cases. In the staple group, the mean age was 58.5 years and the average follow-up was 4.3 years. In the circular plate group, the mean age was 58.6 years and the average follow-up was 3.1 years. Pain, range of motion, grip strength, functional scores (QuickDASH and PWRE), fusion of the midcarpal joint, complications (implant fracture and reoperation) and patients' satisfaction were used as outcome measures. There was no pain in 43% of patients in the staple group and 40% of patients in the circular plate group at the follow-up; range of motion and functional scores were similar in both groups. Seventy-five percent of patients in the staple group were satisfied or very satisfied versus 60% in the circular plate group. The implant broke in 24.3% of cases in the staple group and 60% in the circular plate group. Reoperation was needed in 18% of the staple cases and 14% of the plate cases. There was no difference between the implants in terms of pain, range of motion, functional scores and patient satisfactory. The implant fracture rate in the plate group was high. This study brings into question implant reliability for the four-corner fusion procedure.