Estructura de las comunidades de briófitos en los complejos de páramos de Boyacá-Colombia / Structure of bryophyte communities in the paramo complexes of Boyacá-Colombia

Introducción: Los páramos de Boyacá cubren el 18.3 % de la superficie de Colombia, y son diversos en flora y fauna, además, registran una alta variabilidad climática, topográfica y de hábitats, que permite que estos ecosistemas sean centros de diversidad en el Neotrópico, y por tanto albergan una alta diversidad de briófitos. Objetivo: Analizar la estructura y composición de las comunidades de briófitos de los complejos de páramos de Boyacá. Métodos: a partir de información de literatura, bases de datos y revisión de herbarios, se evaluó la composición florística y la completitud de muestreo para los complejos de páramos y sustratos. Resultados: Se encontraron 5 132 ejemplares, con 343 especies de musgos que fue el grupo más diverso, 256 hepáticas y dos antocerotes. El análisis de completitud de muestreo es representativo en un 98 %. Además, encontramos que la preferencia de sustratos es el terrícola con 409 especies y el cortícola con 341. La diversidad alfa del orden 0D mostró que Tota-Bijagual-Mamapacha (TBM) es el complejo más diverso con 368 especies, y Pisba (124) el menos diverso; el índice 1D mostró que el complejo (TBM) presentó 178 especies consideradas comunes, y la dominancia de especies (2D) fue mayor en el complejo Iguaque-Merchán con 119 taxa dominantes y en menor número Guantiva-La Rusia (105) y TBM (102). Conclusiones: El análisis de la diversidad beta mostró que el 62 % de la disimilitud en la composición de especies entre los complejos se debe al recambio de especies, igualmente sucede con la divergencia por sustratos que es del 51 %. Los briófitos en los páramos de Boyacá representan el 36.05 % de la diversidad colombiana, y el 2.96 % a nivel mundial.

Introduction: The Boyacá paramos cover 18.3 % of the Colombian surface, and are diverse in flora and fauna, moreover, have a high temperature, topography, and habitats, which allow these ecosystems to be centers of diversity in the Neotropics, and therefore they harbor a high diversity of bryophytes. Objectives: Analyze the structure and composition of the bryophyte communities of the paramos in the Boyacá complexes. Methods: Using literature, databases and herbariums records, the floristic composition, and the completeness of the sampling for the paramo and substrate complexes was evaluated. Results: We analyzed 5 132 specimens, with 343 species of mosses being the most diverse group, 256 liverworts and two hornworts. The sampling completeness analysis is 98 % representative. In addition, we found that the preference of substrates is terrestrial with 409 species and corticolous with 341. Alpha diversity of order 0D showed that Tota-Bijagual-Mamapacha (TBM) is the most diverse complex with 368 species, and Pisba (124) the least diverse; the 1D index showed that the complex (TBM) presented 178 species considered common, and the dominance of species (2D) was higher in the Iguaque-Merchán Complex with 119 dominant taxa and Guantiva-La Russia (105) and TBM (105) to a lesser number. (102). Conclusions: the analysis of beta diversity showed that 62 % of the dissimilarity in the composition of species between the complexes is due to the species turnover, the same happens with the divergence by substrates that is 51 %. Bryophytes in the Boyacá paramos represent 36.05 % of Colombian diversity, and 2.96 % worldwide.

Relationships among sporophytic and gametophytic traits of 27 subtropical montane moss species.

PREMISE: Moss sporophytes differ strongly in size and biomass partitioning, potentially reflecting reproductive and dispersal strategies. Understanding how sporophyte traits are coordinated is essential for understanding moss functioning and evolution. This study aimed to answer: (1) how the size and proportions of the sporophyte differ between moss species with and without a prominent central strand in the seta, (2) how anatomical and morphological traits of the seta are related, and (3) how sporophytic biomass relates to gametophytic biomass and nutrient concentrations. METHODS: We studied the relationships between seta anatomical and morphological traits, the biomass of seta, capsule, and gametophyte, and carbon, nitrogen, and phosphorus concentrations of 27 subtropical montane moss species. RESULTS: (1) Moss species with a prominent central strand in the seta had larger setae and heavier capsules than those without a prominent strand. (2) With increasing seta length, setae became thicker and more rounded for both groups, while in species with a prominent central strand, the ratio of transport-cell area to epidermal area decreased. (3) In both groups, mosses with greater gametophytic biomass tended to have heavier sporophytes, but nitrogen and phosphorus concentrations in the gametophyte were unrelated to sporophytic traits. CONCLUSIONS: Our study highlights that the central strand in the seta may have an important functional role and affect the allometry of moss sporophytes. The coordinated variations in sporophyte morphological and anatomical traits follow basic biomechanical principles of cylinder-like structures, and these traits relate only weakly to the gametophytic nutrient concentrations. Research on moss sporophyte functional traits and their relationships to gametophytes is still in its infancy but could provide important insights into their adaptative strategies.

The phylogeny of Syntrichia: An ecologically diverse clade of mosses with an origin in South America.

PREMISE: To address the biodiversity crisis, we need to understand the evolution of all organisms and how they fill geographic and ecological space. Syntrichia is one of the most diverse and dominant genera of mosses, ranging from alpine habitats to desert biocrusts, yet its evolutionary history remains unclear. METHODS: We present a comprehensive phylogenetic analysis of Syntrichia, based on both molecular and morphological data, with most of the named species and closest outgroups represented. In addition, we provide ancestral-state reconstructions of water-related traits and a global biogeographic analysis. RESULTS: We found 10 major well-resolved subclades of Syntrichia that possess geographical or morphological coherence, in some cases representing previously accepted genera. We infer that the extant species diversity of Syntrichia likely originated in South America in the early Eocene (56.5-43.8 million years ago [Mya]), subsequently expanded its distribution to the neotropics, and finally dispersed to the northern hemisphere. There, the clade experienced a recent diversification (15-12 Mya) into a broad set of ecological niches (e.g., the S. caninervis and S. ruralis complexes). The transition from terricolous to either saxicolous or epiphytic habitats occurred more than once and was associated with changes in water-related traits. CONCLUSIONS: Our study provides a framework for understanding the evolutionary history of Syntrichia through the combination of morphological and molecular characters, revealing that migration events that shaped the current distribution of the clade have implications for morphological character evolution in relation to niche diversity.

Microfluidics-Based Bioassays and Imaging of Plant Cells.

Many plant processes occur in the context of and in interaction with a surrounding matrix such as soil (e.g. root growth and root-microbe interactions) or surrounding tissues (e.g. pollen tube growth through the pistil), making it difficult to study them with high-resolution optical microscopy. Over the past decade, microfabrication techniques have been developed to produce experimental systems that allow researchers to examine cell behavior in microstructured environments that mimic geometrical, physical and/or chemical aspects of the natural growth matrices and that cannot be generated using traditional agar plate assays. These microfabricated environments offer considerable design flexibility as well as the transparency required for high-resolution, light-based microscopy. In addition, microfluidic platforms have been used for various types of bioassays, including cellular force assays, chemoattraction assays and electrotropism assays. Here, we review the recent use of microfluidic devices to study plant cells and organs, including plant roots, root hairs, moss protonemata and pollen tubes. The increasing adoption of microfabrication techniques by the plant science community may transform our approaches to investigating how individual plant cells sense and respond to changes in the physical and chemical environment.

Reassessing the role of morphology in bryophyte phylogenetics: Combined data improves phylogenetic inference despite character conflict.

Morphological data has gained renewed attention and has been shown to be crucial in clarifying the phylogenetic relationship in a wide range of taxa. In the last decades, phylogenetic analyses of sequence-level data have radically modified the systematic schemes within bryophytes (early non-vascular land plants) and have revealed a widespread pattern of conflict with morphology-based classifications. Yet, a comprehensive evaluation of character conflict has not yet been performed in the context of combined matrices. In this study, we evaluate the impact of morphology on bryophyte phylogeny following a total-evidence approach across 10 published matrices. The analysed matrices span a wide range of bryophytes, taxonomic levels, gene sampling and number of morphological characters and taxa. Data conflict was addressed by measuring: (i) the topological congruence between individual partitions, (ii) changes in support values of the combined data relative to the molecular partition and (iii) clade stability. The association between these measures and the number of morphological characters per taxon (Nc/T ratio) and the proportion of non-fixed characters (i.e., inapplicable, polymorphic and missing data) was explored. In the individual partition analyses, the Nc/T ratio correlated positively with the topological congruence in six to seven datasets depending on the weighting scheme. The proportion of non-fixed cells had a minor influence on congruence between data partitions. The number of characters and proportion of non-fixed data varied significantly between morphological datasets that improved congruence between data types. This variation suggests that morphological datasets affect the results of combined analyses in different ways, depending on the taxa studied. Combined analyses revealed that, despite the low congruence values between partitions, integrating data types improves support values and stability. However, while non-fixed data had no negative effect on support values, stability was reduced as the proportion of non-fixed cells increased. Nc/T ratio was negatively associated with support values and it showed ambiguous responses in stability evaluations. Overall, the results indicate that adding morphology may contribute to the inference of phylogenetic relationships of bryophytes despite character conflict. Our findings suggest that merely comparing (a) morphology-based classifications with molecular phylogenies or (b) the outcome from individual data partitions can misestimate data conflict. These findings imply that analyses of combined data may provide conservative assessments of data conflict and, eventually, lead to an improved sampling of morphological characters in large-scale analyses of bryophytes.

Canopy leaf area index at its higher end: dissection of structural controls from leaf to canopy scales in bryophytes.

There is evidence that mosses with miniature foliage elements have extremely large leaf area index (LAI) values, but it is unclear what canopy traits are responsible for these high LAI values in architecturally divergent mosses, and how the inherent trade-offs limiting maximum LAI in vascular plants can be overcome in mosses. To determine the quantitative significance of different traits in determining LAI, we developed a method to dissect LAI into underlying functionally dependent constituent traits at leaf, shoot and canopy scales. The suites of structural traits were studied altogether for 43 moss canopies from 11 species with contrasting light and water requirements along gap-understory gradients to obtain as large a range of variation in moss architecture as possible and evaluate the differentiation in moss LAI in relation to species ecology. Extensive variation in moss structural traits, 11- (shoot length) to 77-fold (shoot number per area, NS¯ ), was observed at all structural scales from leaf to canopy. However, LAI only varied nine-fold, as the result of two key trade-offs: leaf size vs number trade-off and shoot leaf area vs shoot density trade-off. Owing to these negative relationships, and greater variability in NS¯ , LAI primarily scaled with NS¯ . NS¯ and LAI increased with site light availability, and LAI was greater in open and dry habitat species. This study highlights a huge structural diversity among moss canopies, but indicates that canopies converge to a much narrower range of LAI due to trait trade-offs such that, counterintuitively, minute leaf size and densely leafed stems are not necessarily responsible for high LAI in mosses.

Anatomical constraints to nonstomatal diffusion conductance and photosynthesis in lycophytes and bryophytes.

Photosynthesis in bryophytes and lycophytes has received less attention than terrestrial plant groups. In particular, few studies have addressed the nonstomatal diffusion conductance to CO2 gnsd of these plant groups. Their lower photosynthetic rate per leaf mass area at any given nitrogen concentration compared with vascular plants suggested a stronger limitation by CO2 diffusion. We hypothesized that bryophyte and lycophyte photosynthesis is largely limited by low gnsd . Here, we studied CO2 diffusion inside the photosynthetic tissues and its relationships with photosynthesis and anatomical parameters in bryophyte and lycophyte species in Antarctica, Australia, Estonia, Hawaii and Spain. On average, lycophytes and, specially, bryophytes had the lowest photosynthetic rates and nonstomatal diffusion conductance reported for terrestrial plants. These low values are related to their very thick cell walls and their low exposure of chloroplasts to cell perimeter. We conclude that the reason why bryophytes lie at the lower end of the leaf economics spectrum is their strong nonstomatal diffusion conductance limitation to photosynthesis, which is driven by their specific anatomical characteristics.

Contributions to the bryological knowledge of ASPA 125, Fildes Peninsula, King George Island.

BACKGROUND: With 29 Km2, the Fildes Peninsula is the largest ice free area in King George Island and probably in Antarctica. The region is house of six permanent bases including the only airport in the South Shetlands, which led to impacts on its original landscape and vegetation. In recognition for the need to protect natural values, an Antarctic Specially Protected Area (ASPA 125) was established in the region. Focused mostly on protecting the fossils, the ASPA also plays a role in protecting the vegetation but so far, the management plan for the area does not contain a list of moss species present there. RESULTS: We provided an updated study and checklist of mosses present in ASPA 125. A key to species identification and photographs of main morphological features are also available in this paper. Also, six new occurrences are reported for Fildes Peninsula. CONCLUSION: Considering the scarce knowledge about specific local floras in Antarctica associated with highly impacted area, of which only a fraction is protected, it is suggested the necessity to invest in detailed sampling studies, as well as in a better understanding of the local floras interactions in Antarctica.

Canopy structure affects temperature distributions and free convection in moss shoot systems.

PREMISE OF THE STUDY: Nonvascular plants play important roles in exchange of water and heat at the soil-atmosphere interface. Differential evaporative cooling may cause temperature gradients within bryophyte canopies, influencing convective heat and mass transport. Understanding mechanisms that affect fluxes through moss layers should improve models of forest floor function. METHODS: A three-dimensional thermal imaging system measured temperature distributions within moss shoot systems that were used to explore relationships among canopy structure, temperature gradients, evaporation, and conductance to water vapor (gs ). We studied five moss species under dark and light conditions in the lab. Also, these properties were measured in two species that differed in canopy structure during drying. KEY RESULTS: Differential evaporative cooling led to a 1.4 to 5.0°C range in shoot temperatures within canopies. Samples displayed -0.5 to -0.9°C/cm temperature gradients with cooler apical temperatures. Gradient magnitudes did not differ among species, but taller canopies expressed greater temperature differences. Light enhanced both the gradient and the temperature difference. Rates of evaporation were significantly related to canopy height in the light, but not in the dark, although gs was positively associated with canopy height in both. Rayleigh (Ra) numbers characterize whether temperature gradients likely generate free convection. In tall canopies, Ra numbers exceeded the value indicative of free convection and turbulent flow. As plants dried, temperature gradients decreased. CONCLUSIONS: When moss canopies are wet, cooler apical temperatures create thermal instabilities within the canopies that appear sufficient to enhance convective transport of water vapor and heat in tall canopies with low bulk density.

Wanted dead or alive (probably dead): Stem group Polytrichaceae.

PREMISE OF THE STUDY: The Polytrichaceae are a widespread and morphologically isolated moss lineage. Early attempts to characterize phylogenetic relationships within the family suggested that morphology is not phylogenetically informative. Two well-characterized fossils similar to basal and derived Polytrichaceae (Meantoinea alophosioides and Eopolytrichum antiquum, respectively), are known from Cretaceous rocks. To assess the phylogenetic positions of these fossils and compare hypotheses of relationships recovered using molecular vs. morphological methods, we conducted a comprehensive morphology-based phylogenetic study of Polytrichaceae. METHODS: We evaluated the phylogenetic relationships of Polytrichaceae using a data set of 100 morphological characters (including 11 continuously varying traits codified as continuous characters) scored for 44 species of acrocarpous mosses and parsimony as the optimality criterion. KEY RESULTS: Continuous characters significantly increased the resolving power of the analyses. The overall ingroup topology was sensitive to rooting as determined by outgroup selection, with some analyses yielding results that were incongruent with those of molecular studies. Both fossils had stable phylogenetic relationships, irrespective of outgroup sampling. CONCLUSIONS: Our results suggest that morphology is useful in resolving phylogenetic relationships in the Polytrichaceae, if both discrete and continuous characters are used. However, our rooting experiments demonstrate that there is no superior way to root analyses and indicate that relationships within the family are best evaluated using unrooted networks without outgroup taxa. These rooting problems suggest that additional information is needed to understand the phylogenetic relationships of Polytrichaceae. Such additional information could come from fossils of stem group polytrichaceous mosses, which await discovery.

Composición y distribución de hepáticas (Marchantiophyta) en un intervalo altitudinal en la Cordillera Oriental de Colombia / Composition and distribution of liverworts (Marchantiophyta) in a continuous altitudinal range on the Cordillera Oriental of Colombia

Resumen La composición de especies de hepáticas sobre los Andes del Norte varía con la altitud en relación con diferentes factores bióticos (vegetación) y abióticos (temperatura, humedad y luminosidad). Para probar esta afirmación se determinó la diversidad y distribución de las Marchantiophytas en un rango altitudinal entre los 2 400 y 3 400 m sobre la Cordillera Oriental colombiana, estableciendo cómo éstas varían entre altitudes y zonas de vida con respecto a los factores abióticos. Se realizaron muestreos cada 200 m y se registraron datos ambientales durante un año. Se encontraron 162 especies, la composición de hepáticas varió a lo largo del gradiente altitudinal con un gran número de especies únicas en cada altitud. Se encontró la máxima diversidad en los 3 000 m, con una dominancia del hábito folioso y de especies epifitas, mientras que la mayor riqueza de familias y géneros se presentó entre los 3 200 y 3 400 m. Anoplolejeunea conferta presentó la mayor cobertura en la zona así como el mayor valor del índice de valor de importancia (IVI) y distribución altitudinal más amplia, ubicándose desde los 2 400 hasta 3 000 m en cuatro diferentes sustratos. Las hepáticas se distribuyeron altitudinalmente de manera diferencial, con una diversidad beta alta (0.864) debido al recambio altitudinal de especies, con más del 30 % de disimilaría en la composición de especies cada 200 m altitudinales, dependiendo principalmente de cambios en la luminosidad. Consecuentemente, la temperatura, humedad y luminosidad y su relación con la vegetación son determinantes en la diversidad y distribución de las hepáticas en la Cordillera Oriental de los Andes colombianos.

Abstract The species composition of liverworts varies with altitude in the Northern part of the Andes due to the relationship of biotic (vegetation) and abiotic factors (temperature, humidity, and sunlight brightness). In order to test this affirmation we determined the diversity, species composition, abundance, and distribution of species of Marchantiophyta in an altitudinal range from 2 400 to 3 400 m on the Colombian Cordillera Oriental, and established how these vary with altitude and life-zones, regarding abiotic factors. Samples, taken every 200 m in the altitudinal range, and environmental data were registered during a year. We found 162 species, the composition of liverworts varied throughout the altitudinal gradient with a high number of unique species in each altitude. The maximum diversity was found at 3 000 m, along with a dominance of leafy habit and epiphytic species, while the higher richness of families and genera was found between 3 200 and 3 400 m. Anoplolejeunea conferta had the largest value of coverage in the zone, the highest value of importance value index (IVI), and the widest altitudinal distribution, from 2 400 to 3 000 m in four different substrates. The liverworts were distributed differently with altitude, we found a high beta diversity (0.864) due to the replacement of species, with more that 30 % of dissimilarity in species composition every 200 altitudinal meters, mainly depending on sunlight brightness variation. Accordingly, we found that abiotic factors like temperature, humidity, and sunlight brightness and its relationship vegetation are determinant in the diversity and altitudinal distribution of liverworts in the study area. Rev. Biol. Trop. 66(2): 559-570. Epub 2018 June 01.

Development of the sphagnoid areolation pattern in leaves of Palaeozoic protosphagnalean mosses.

Background and Aims: Protosphagnalean mosses constitute the largest group of extinct mosses of still uncertain affinity. Having the general morphology of the Bryopsida, some have leaves with an areolation pattern characteristic of modern Sphagna. This study describes the structure and variation of these patterns in protosphagnalean mosses and provides a comparison with those of modern Sphagna. Methods: Preparations of fossil mosses showing preserved leaf cell structure were obtained by dissolving rock, photographed, and the resulting images were transformed to graphical format and analysed with Areoana computer software. Key Results: The sphagnoid areolation pattern is identical in its basic structure for both modern Sphagnum and Palaeozoic protosphagnalean mosses. However, in the former group the pattern develops through unequal oblique cell divisions, while in the latter the same pattern is a result of equal cell divisions taking place in a specific order with subsequent uneven cell growth. The protosphagnalean pathway leads to considerable variability in leaf structure. Conclusions: Protosphagnalean mosses had a unique ability to switch the development of leaf areolation between a pathway unique to Sphagnum and another one common to all other mosses. This developmental polyvariancy hinders attempts to classify these mosses, as characters previously considered to be of generic significance can be shown to co-occur in one individual leaf. New understanding of the ontogeny has allowed us to re-evaluate the systematic significance of such diagnostic characters in these Palaeozoic plants, showing that their similarity to Sphagnum is less substantial.

Grimmiaceae in the Early Cretaceous: Tricarinella crassiphylla gen. et sp. nov. and the value of anatomically preserved bryophytes.

Background and Aims: Widespread and diverse in modern ecosystems, mosses are rare in the fossil record, especially in pre-Cenozoic rocks. Furthermore, most pre-Cenozoic mosses are known from compression fossils, which lack detailed anatomical information. When preserved, anatomy significantly improves resolution in the systematic placement of fossils. Lower Cretaceous (Valanginian) deposits on Vancouver Island (British Columbia, Canada) contain a diverse anatomically preserved flora including numerous bryophytes, many of which have yet to be characterized. Among them is the grimmiaceous moss described here. Methods: One fossil moss gametophyte preserved in a carbonate concretion was studied in serial sections prepared using the cellulose acetate peel technique. Key Results: Tricarinella crassiphylla gen. et sp. nov. is a moss with tristichous phyllotaxis and strongly keeled leaves. The combination of an acrocarpous condition (inferred based on a series of morphological features), a central conducting strand, a homogeneous leaf costa and a lamina with bistratose portions and sinuous cells, and multicellular gemmae, supports placement of Tricarinella in family Grimmiaceae. Tricarinella is similar to Grimmia, a genus that exhibits broad morphological variability. However, tristichous phyllotaxis and especially the lamina, bistratose at the base but not in distal portions of the leaf, set Tricarinella apart as a distinct genus. Conclusions: Tricarinella crassiphylla marks the oldest record for both family Grimmiaceae and sub-class Dicranidae, providing a hard minimum age (136 million years) for these groups. The fact that this fossil could be placed in an extant family, despite a diminutive size, emphasizes the considerable resolving power of anatomically preserved bryophyte fossils, even when recovered from allochthonous assemblages of marine sediments, such as the Apple Bay flora. Discovery of Tricarinella re-emphasizes the importance of paleobotanical studies as the only approach allowing access to a significant segment of biodiversity, the extinct biodiversity, which is unattainable by other means of investigation.

Contributions to the bryological knowledge of ASPA 125, Fildes Peninsula, King George Island

BACKGROUND: With 29 Km2, the Fildes Peninsula is the largest ice free area in King George Island and probably in Antarctica. The region is house of six permanent bases including the only airport in the South Shetlands, which led to impacts on its original landscape and vegetation. In recognition for the need to protect natural values, an Antarctic Specially Protected Area (ASPA 125) was established in the region. Focused mostly on protecting the fossils, the ASPA also plays a role in protecting the vegetation but so far, the management plan for the area does not contain a list of moss species present there. RESULTS: We provided an updated study and checklist of mosses present in ASPA 125. A key to species identification and photographs of main morphological features are also available in this paper. Also, six new occurrences are reported for Fildes Peninsula. CONCLUSION: Considering the scarce knowledge about specific local floras in Antarctica associated with highly impacted area, of which only a fraction is protected, it is suggested the necessity to invest in detailed sampling studies, as well as in a better understanding of the local floras interactions in Antarctica.

Multiple innovations underpinned branching form diversification in mosses.

Broad-scale evolutionary comparisons have shown that branching forms arose by convergence in vascular plants and bryophytes, but the trajectory of branching form diversification in bryophytes is unclear. Mosses are the most species-rich bryophyte lineage and two sub-groups are circumscribed by alternative reproductive organ placements. In one, reproductive organs form apically, terminating growth of the primary shoot (gametophore) axis. In the other, reproductive organs develop on very short lateral branches. A switch from apical to lateral reproductive organ development is proposed to have primed branching form diversification. Moss gametophores have modular development and each module develops from a single apical cell. Here we define the architectures of 175 mosses by the number of module classes, branching patterns and the pattern in which similar modules repeat. Using ancestral character state reconstruction we identify two stages of architectural diversification. During a first stage there were sequential changes in the module repetition pattern, reproductive organ position, branching pattern and the number of module classes. During a second stage, vegetative changes occurred independently of reproductive fate. The results pinpoint the nature of developmental change priming branching form diversification in mosses and provide a framework for mechanistic studies of architectural diversification.

Extending the fossil record of Polytrichaceae: Early Cretaceous Meantoinea alophosioides gen. et sp. nov., permineralized gametophytes with gemma cups from Vancouver Island.

PREMISE OF THE STUDY: Diverse in modern ecosystems, mosses are dramatically underrepresented in the fossil record. Furthermore, most pre-Cenozoic mosses are known only from compression fossils, lacking detailed anatomical information. When preserved, anatomy vastly improves resolution in the systematic placement of fossils. Lower Cretaceous deposits at Apple Bay (Vancouver Island, British Columbia, Canada) contain a diverse anatomically preserved flora that includes numerous bryophytes, many of which have yet to be characterized. Among them is a polytrichaceous moss that is described here. METHODS: Fossil moss gametophytes preserved in four carbonate concretions were studied in serial sections prepared using the cellulose acetate peel technique. KEY RESULTS: We describe Meantoinea alophosioides gen. et sp. nov., a polytrichaceous moss with terminal gemma cups containing stalked, lenticular gemmae. Leaves with characteristic costal anatomy, differentiated into sheathing base and free lamina and bearing photosynthetic lamellae, along with a conducting strand in the stem, place Meantoinea in family Polytrichaceae. The bistratose leaf lamina with an adaxial layer of mamillose cells, short photosynthetic lamellae restricted to the costa, and presence of gemma cups indicate affinities with basal members of the Polytrichaceae, such as Lyellia, Bartramiopsis, and Alophosia. CONCLUSIONS: Meantoinea alophosioides enriches the documented moss diversity of an already-diverse Early Cretaceous plant fossil assemblage. This is the third moss described from the Apple Bay plant fossil assemblage and represents the first occurrence of gemma cups in a fossil moss. It is also the oldest unequivocal record of Polytrichaceae, providing a hard minimum age for the group of 136 million years.

On the benefits of living in clumps: a case study on Polytrichastrum formosum.

The study concerns the mechanics and water relationships of clumps of a species of endohydric moss, Polytrichastrum formosum. Anatomical and morphological studies were done using optical and scanning electron microscopy. Experiments on waterdrop capture and their distribution to adjacent shoots within a moss clump were performed with the experimental set-up for the droplet collision phenomena and ultra-high speed camera. The mechanical strength of the moss clump was tested on an electromechanical testing machine. During the process of moss clump wetting, the falling water drops were captured by the apical stem part or leaves, then flowed down while adhering to the gametophore and never lost their surface continuity. In places of contact with another leaf, the water drop stops there and joins the leaves, enabling their hydration. Mathematical analysis of anatomical images showed that moss stems have different zones with varying cell lumen and cell wall/cell radius ratios, suggesting the occurrence of a periodic component structure. Our study provides evidence that the reaction of mosses to mechanical forces depends on the size of the clump, and that small groups are clearly stronger than larger groups. The clump structure of mosses acts as a net for falling rain droplets. Clumps of Polytrichastrum having overlapping leaves, at the time of loading formed a structure similar to a lattice. The observed reaction of mosses to mechanical forces indicates that this phenomenon appears to be analogous to the 'size effect on structural strength' that is of great importance for various fields of engineering.

Dehydration-responsive features of Atrichum undulatum.

Drought is an increasingly important limitation on plant productivity worldwide. Understanding the mechanisms of drought tolerance in plants can lead to new strategies for developing drought-tolerant crops. Many moss species are able to survive desiccation-a more severe state of dehydration than drought. Research into the mechanisms and evolution of desiccation tolerance in basal land plants is of particular significance to both biology and agriculture. In this study, we conducted morphological, cytological, and physiological analyses of gametophytes of the highly desiccation-tolerant bryophyte Atrichum undulatum (Hedw.) P. Beauv during dehydration and rehydration. Our results suggested that the mechanisms underlying the dehydration-recovery cycle in A. undulatum gametophytes include maintenance of membrane stability, cellular structure protection, prevention of reactive oxygen species (ROS) generation, elimination of ROS, protection against ROS-induced damage, and repair of ROS-induced damage. Our data also indicate that this dehydration-recovery cycle consists not only of the physical removal and addition of water, but also involves a highly organized series of cytological, physiological, and biochemical changes. These attributes are similar to those reported for other drought- and desiccation-tolerant plant species. Our findings provide major insights into the mechanisms of dehydration-tolerance in the moss A. undulatum.

Crown Group Lejeuneaceae and Pleurocarpous Mosses in Early Eocene (Ypresian) Indian Amber.

Cambay amber originates from the warmest period of the Eocene, which is also well known for the appearance of early angiosperm-dominated megathermal forests. The humid climate of these forests may have triggered the evolution of epiphytic lineages of bryophytes; however, early Eocene fossils of bryophytes are rare. Here, we present evidence for lejeuneoid liverworts and pleurocarpous mosses in Cambay amber. The preserved morphology of the moss fossil is inconclusive for a detailed taxonomic treatment. The liverwort fossil is, however, distinctive; its zig-zagged stems, suberect complicate-bilobed leaves, large leaf lobules, and small, deeply bifid underleaves suggest a member of Lejeuneaceae subtribe Lejeuneinae (Harpalejeunea, Lejeunea, Microlejeunea). We tested alternative classification possibilities by conducting divergence time estimates based on DNA sequence variation of Lejeuneinae using the age of the fossil for corresponding age constraints. Consideration of the fossil as a stem group member of Microlejeunea or Lejeunea resulted in an Eocene to Late Cretaceous age of the Lejeuneinae crown group. This reconstruction is in good accordance with published divergence time estimates generated without the newly presented fossil evidence. Balancing available evidence, we describe the liverwort fossil as the extinct species Microlejeunea nyiahae, representing the oldest crown group fossil of Lejeuneaceae.

Endemism in the moss flora of North America.

PREMISE OF THE STUDY: Identifying regions of high endemism is a critical step toward understanding the mechanisms underlying diversification and establishing conservation priorities. Here, we identified regions of high moss endemism across North America. We also identified lineages that contribute disproportionately to endemism and document the progress of efforts to inventory the endemic flora. METHODS: To understand the documentation of endemic moss diversity in North America, we tabulated species publication dates to document the progress of species discovery across the continent. We analyzed herbarium specimen data and distribution data from the Flora of North America project to delineate major regions of moss endemism. Finally, we surveyed the literature to assess the importance of intercontinental vs. within-continent diversification for generating endemic species. KEY RESULTS: Three primary regions of endemism were identified and two of these were further divided into a total of nine subregions. Overall endemic richness has two peaks, one in northern California and the Pacific Northwest, and the other in the southern Appalachians. Description of new endemic species has risen steeply over the last few decades, especially in western North America. Among the few studies documenting sister species relationships of endemics, recent diversification appears to have played a larger role in western North America, than in the east. CONCLUSIONS: Our understanding of bryophyte endemism continues to grow rapidly. Large continent-wide data sets confirm early views on hotspots of endemic bryophyte richness and indicate a high rate of ongoing species discovery in North America.