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Primary producers shape terrestrial biodiversity, but most research has focused on vascular plants, while the role of cryptogams (mosses, lichens and algae) remains under‐explored. Cryptogams dominate Antarctic vegetation and support diverse microarthropod communities. However, how cryptogam traits influence these communities remains poorly understood. We therefore investigated the role of 28 cryptogam species and one vascular plant, via their functional traits, in shaping microarthropod communities across three contrasting sites (Signy Island, Byers Peninsula and Rothera) in the maritime Antarctic. We hypothesized that vegetation nitrogen and moisture content, major microarthropod taxa, and abiotic drivers interact to influence community patterns. Vegetation type effects on total microarthropod abundance were context‐dependent. Mosses hosted more microarthropods than lichens at Signy Island, but these differences diminished further south. Microarthropod richness and springtail abundance were consistently higher in mosses than lichens across all sites, whereas mite abundance did not differ between vegetation types. Cryptogam nitrogen and moisture content strongly predicted microarthropod community patterns, although their influence varied with vegetation type and location. Among mosses, moisture increased springtail abundance but reduced diversity due to the dominance of Cryptopygus antarcticus. In lichens, nitrogen had a stronger influence than in mosses, particularly on mite abundance and Shannon diversity. As hypothesized, moisture was more important at the harshest southern site, while nitrogen had stronger effects at more productive northern locations. These findings emphasize that the influence of cryptogam traits in structuring Antarctic terrestrial biodiversity is modulated by the environmental context. With future shifts predicted in vegetation composition, the functional traits of emerging dominant species may restructure microarthropod communities and their ecological functions.

DNA barcoding involves sequencing a standard region of DNA as a tool for species identification. However, there has been no agreement on which region(s) should be used for barcoding land plants. To provide a community recommendation on a standard plant barcode, we have compared the performance of 7 leading candidate plastid DNA regions (atpF-atpH spacer, matK gene, rbcL gene, rpoB gene, rpoC1 gene, psbK-psbI spacer, and trnH-psbA spacer). Based on assessments of recoverability, sequence quality, and levels of species discrimination, we recommend the 2-locus combination of rbcL+matK as the plant barcode. This core 2-locus barcode will provide a universal framework for the routine use of DNA sequence data to identify specimens and contribute toward the discovery of overlooked species of land plants.

• Trees are known to emit methane (CH₄) and nitrous oxide (N₂O), with tropical wetland trees being considerable CH₄ sources. Little is known about CH₄ and especially N₂O exchange of trees growing in tropical rain forests under nonflooded conditions. • We determined CH₄ and N₂O exchange of stems of six dominant tree species, cryptogamic stem covers, soils and volcanic surfaces at the start of the rainy season in a 400-yr-old tropical lowland rain forest situated on a basaltic lava flow (Réunion Island). We aimed to understand the unknown role in greenhouse gas fluxes of these atypical tropical rain forests on basaltic lava flows. • The stems studied were net sinks for atmospheric CH₄ and N₂O, as were cryptogams, which seemed to be co-responsible for the stem uptake. In contrast with more commonly studied rain forests, the soil and previously unexplored volcanic surfaces consumed CH₄. Their N₂O fluxes were negligible. • Greenhouse gas uptake potential by trees and cryptogams constitutes a novel and unique finding, thus showing that plants can serve not only as emitters, but also as consumers of CH₄ and N₂O. The volcanic tropical lowland rain forest appears to be an important CH₄ sink, as well as a possible N₂O sink.

The increase in aridity predicted with climate change will have a negative impact on the multiple functions and services (multifunctionality) provided by dryland ecosystems worldwide. In these ecosystems, soil communities dominated by mosses, lichens and cyanobacteria (biocrusts) play a key role in supporting multifunctionality. However, whether biocrusts can buffer the negative impacts of aridity on important biogeochemical processes controlling carbon (C), nitrogen (N), and phosphorus (P) pools and fluxes remains largely unknown. Here, we conducted an empirical study, using samples from three continents (North America, Europe and Australia), to evaluate how the increase in aridity predicted by climate change will alter the capacity of biocrust-forming mosses to modulate multiple ecosystem processes related to C, N and P cycles. Compared with soil surfaces lacking biocrusts, biocrust-forming mosses enhanced multiple functions related to C, N and P cycling and storage in semiarid and arid, but not in humid and dry-subhumid, environments. Most importantly, we found that the relative positive effects of biocrust-forming mosses on multifunctionality compared with bare soil increased with increasing aridity. These results were mediated by plant cover and the positive effects exerted by biocrust-forming mosses on the abundance of soil bacteria and fungi. Our findings provide strong evidence that the maintenance of biocrusts is crucial to buffer negative effects of climate change on multifunctionality in global drylands.

Global cooling and glacial-interglacial cycles since Antarctica’s isolation have been responsible for the diversification of the region’s marine fauna. By contrast, these same Earth system processes are thought to have played little role terrestrially, other than driving widespread extinctions. Here, we show that on islands along the Antarctic Polar Front, paleoclimatic processes have been key to diversification of one of the world’s most geographically isolated and unique groups of herbivorous beetles-Ectemnorhinini weevils. Combining phylogenomic, phylogenetic, and phylogeographic approaches, we demonstrate that these weevils colonized the sub-Antarctic islands from Africa at least 50 Ma ago and repeatedly dispersed among them. As the climate cooled from the mid-Miocene, diversification of the beetles accelerated, resulting in two species-rich clades. One of these clades specialized to feed on cryptogams, typical of the polar habitats that came to prevail under Miocene conditions yet remarkable as a food source for any beetle. This clade’s most unusual representative is a marine weevil currently undergoing further speciation. The other clade retained the more common weevil habit of feeding on angiosperms, which likely survived glaciation in isolated refugia. Diversification of Ectemnorhinini weevils occurred in synchrony with many other Antarctic radiations, including penguins and notothenioid fishes, and coincided with major environmental changes. Our results thus indicate that geo-climatically driven diversification has progressed similarly for Antarctic marine and terrestrial organisms since the Miocene, potentially constituting a general biodiversity paradigm that should be sought broadly for the region’s taxa.

BACKGROUND: Recent decades have seen a major surge in the study of interspecific variation in functional traits in comparative plant ecology, as a tool to understanding and predicting ecosystem functions and their responses to environmental change. However, this research has been biased almost exclusively towards vascular plants. Very little is known about the role and applicability of functional traits of non-vascular cryptogams, particularly bryophytes and lichens, with respect to biogeochemical cycling. Yet these organisms are paramount determinants of biogeochemistry in several biomes, particularly cold biomes and tropical rainforests, where they: (1) contribute substantially to above-ground biomass (lichens, bryophytes); (2) host nitrogen-fixing bacteria, providing major soil N input (lichens, bryophytes); (3) control soil chemistry and nutrition through the accumulation of recalcitrant polyphenols (bryophytes) and through their control over soil and vegetation hydrology and temperatures; (4) both promote erosion (rock weathering by lichens) and prevent it (biological crusts in deserts); (5) provide a staple food to mammals such as reindeer (lichens) and arthropodes, with important feedbacks to soils and biota; and (6) both facilitate and compete with vascular plants. APPROACH: Here we review current knowledge about interspecific variation in cryptogam traits with respect to biogeochemical cycling and discuss to what extent traits and measuring protocols needed for bryophytes and lichens correspond with those applied to vascular plants. We also propose and discuss several new or recently introduced traits that may help us understand and predict the control of cryptogams over several aspects of the biogeochemistry of ecosystems. CONCLUSIONS: Whilst many methodological challenges lie ahead, comparative cryptogam ecology has the potential to meet some of the important challenges of understanding and predicting the biogeochemical and climate consequences of large-scale environmental changes driving shifts in the cryptogam components of vegetation composition.

• Primary succession models focus on aboveground vascular plants. However, the prevalence of mosses and lichens, that is cryptogams, suggests they play a role in soil successions. Here, we explore whether effects of cryptogams on belowground microbes can facilitate progressive shifts in sand dune succession. • We linked aboveground vegetation, belowground bacterial and fungal communities, and soil chemical properties in six successional stages in Arctic inland sand dunes: bare sand, grass, moss, lichen, ericoid heath and mountain birch forest. • Compared with the bare sand and grass stages, microbial biomass and the proportion of fungi increased in the moss stage, and later stage microbial groups appeared despite the absence of their host plants. Microbial communities of the lichen stage resembled the communities in the vascular plant stages. Bacterial communities correlated better with soil chemical variables than with vegetation and vice versa for fungal communities. The correlation of fungi with vegetation increased with vascular vegetation. • Distinct bacterial and fungal patterns of biomass, richness and plant–microbe interactions showed that the aboveground vegetation change structured the bacterial and fungal community differently. The asynchrony of aboveground vs belowground changes suggests that cryptogams can drive succession towards vascular plant dominance through microbially mediated facilitation in eroded Arctic soil.

Background and AimsThroughout the history of fern classification, familial and generic concepts have been highly labile. Many classifications and evolutionary schemes have been proposed during the last two centuries, reflecting different interpretations of the available evidence. Knowledge of fern structure and life histories has increased through time, providing more evidence on which to base ideas of possible relationships, and classification has changed accordingly. This paper reviews previous classifications of ferns and presents ideas on how to achieve a more stable consensus.ScopeAn historical overview is provided from the first to the most recent fern classifications, from which conclusions are drawn on past changes and future trends. The problematic concept of family in ferns is discussed, with a particular focus on how this has changed over time. The history of molecular studies and the most recent findings are also presented.Key ResultsFern classification generally shows a trend from highly artificial, based on an interpretation of a few extrinsic characters, via natural classifications derived from a multitude of intrinsic characters, towards more evolutionary circumscriptions of groups that do not in general align well with the distribution of these previously used characters. It also shows a progression from a few broad family concepts to systems that recognized many more narrowly and highly controversially circumscribed families; currently, the number of families recognized is stabilizing somewhere between these extremes. Placement of many genera was uncertain until the arrival of molecular phylogenetics, which has rapidly been improving our understanding of fern relationships. As a collective category, the so-called ‘fern allies’ (e.g. Lycopodiales, Psilotaceae, Equisetaceae) were unsurprisingly found to be polyphyletic, and the term should be abandoned. Lycopodiaceae, Selaginellaceae and Isoëtaceae form a clade (the lycopods) that is sister to all other vascular plants, whereas the whisk ferns (Psilotaceae), often included in the lycopods or believed to be associated with the first vascular plants, are sister to Ophioglossaceae and thus belong to the fern clade. The horsetails (Equisetaceae) are also members of the fern clade (sometimes inappropriately called ‘monilophytes’), but, within that clade, their placement is still uncertain. Leptosporangiate ferns are better understood, although deep relationships within this group are still unresolved. Earlier, almost all leptosporangiate ferns were placed in a single family (Polypodiaceae or Dennstaedtiaceae), but these families have been redefined to narrower more natural entities.ConclusionsConcluding this paper, a classification is presented based on our current understanding of relationships of fern and lycopod clades. Major changes in our understanding of these families are highlighted, illustrating issues of classification in relation to convergent evolution and false homologies. Problems with the current classification and groups that still need study are pointed out. A summary phylogenetic tree is also presented. A new classification in which Aspleniaceae, Cyatheaceae, Polypodiaceae and Schizaeaceae are expanded in comparison with the most recent classifications is presented, which is a modification of those proposed by Smith et al. (2006, 2008) and Christenhusz et al. (2011). These classifications are now finding a wider acceptance and use, and even though a few amendments are made based on recently published results from molecular analyses, we have aimed for a stable family and generic classification of ferns.

We explored the influence of climatic factors on diversity patterns of multiple taxa (lichens, bryophytes, and vascular plants) along a steep elevational gradient to predict communities’ dynamics under future climate change scenarios in Mediterranean regions. We analysed (1) species richness patterns in terms of heat-adapted, intermediate, and cold-adapted species; (2) pairwise beta-diversity patterns, also accounting for its two different components, species replacement and richness difference; (3) the influence of climatic variables on species functional traits. Species richness is influenced by different factors between three taxonomic groups, while beta diversity differs mainly between plants and cryptogams. Functional traits are influenced by different factors in each taxonomic group. On the basis of our observations, poikilohydric cryptogams could be more impacted by climate change than vascular plants. However, contrasting species-climate and traits-climate relationships were also found between lichens and bryophytes suggesting that each group may be sensitive to different components of climate change. Our study supports the usefulness of a multi-taxon approach coupled with a species traits analysis to better unravel the response of terrestrial communities to climate change. This would be especially relevant for lichens and bryophytes, whose response to climate change is still poorly explored.