Trade-off between seed dispersal in space and time.

Seed movement and delayed germination have long been thought to represent alternative risk-spreading strategies, but current evidence covers limited scales and yields mixed results. Here we present the first global-scale test of a negative correlation between dispersal and dormancy. The result demonstrates a strong and consistent pattern that species with dormant seeds have reduced spatial dispersal, also in the context of life-history traits such as seed mass and plant lifespan. Long-lived species are more likely to have large, non-dormant seeds that are dispersed far. Our findings provide robust support for the theoretical prediction of a dispersal trade-off between space and time, implying that a joint consideration of risk-spreading strategies is imperative in studying plant life-history evolution. The bet-hedging patterns in the dispersal-dormancy correlation and the associated reproductive traits have implications for biodiversity conservation, via prediction of which plant groups would be most impacted in the changing era.

Macroevolutionary patterns in seed component mass and different evolutionary trajectories across seed desiccation responses.

Seed coat and seed reserve show substantial mass variation, play different roles in plant life strategies and are shaped by different selective forces. However, remarkably little is known about the macroevolution of the relative allocation in seed components and its influence on important ecophysiological processes. Using phylogenetic comparative methods and evolutionary modelling approaches, we modelled mass changes in seed components along individual lineages for 940 species and compared the patterns across seed desiccation responses. Seed component allocation was driven primarily by changes in reserve mass rather than coat mass, as evolutionary rates in reserve mass significantly outpaced those in coat mass. Although the scaling patterns between reserve mass and coat mass were similar across desiccation responses, desiccation-sensitive seeds allocated more and evolved faster in reserve compared to desiccation-tolerant seeds. The findings emphasize the relative importance of reserve to coat in the evolution of plant reproductive strategies, revealing potential ecological advantages gained by enlarged reserve. As the first quantification of the evolutionary tempo and mode of seed component mass, our study allows a detailed interpretation of evolutionary pathways underlying seed storage behaviours and advances the understanding of the evolution of desiccation sensitivity in seeds.

Taxonomic affinity, habitat and seed mass strongly predict seed desiccation response: a boosted regression trees analysis based on 17 539 species.

Background and Aims: Seed desiccation response plays an important role in plant regeneration ecology, and has significant implications for species conservation. The majority of seed plants produce desiccation-tolerant (orthodox) seeds, whilst comparatively few produce desiccation-sensitive (recalcitrant) seeds that are unable to survive dehydration, and which cannot be conserved in traditional seed banks. This study develops a set of models to predict seed desiccation response in unstudied species. Methods: Taxonomy, trait, location and climate data were compiled to form a global data set of 17 539 species. Three boosted regression trees models were then developed to predict species' seed desiccation responses based on habitat and trait information for the species, and the seed desiccation responses of close relatives (either members of the same genus, family or order, depending on the model). Ten-fold cross-validation was used to test model predictive success. The utility of the models was then demonstrated by predicting seed desiccation response for two floras: Ecuador, and Britain and Ireland. Key Results: The three models had varying success rates for identifying the desiccation-sensitive species: 89 % for the genus-level model, 79 % for the family-level model and 60 % for the order-level model. The most important predictor variables were the seed desiccation responses of a species' relatives, seed mass and annual precipitation. It is predicted that 10 % of seed plants from Ecuador and 1.2 % of those from Britain and Ireland produce desiccation-sensitive seeds. Due to data availability, prediction accuracy is likely to be higher for the British and Irish flora, where it is estimated that a desiccation-sensitive species had a 96.7 % chance of being correctly identified, compared with 80.8 % in the Ecuador flora. Conclusions: These models can utilize existing data to predict species' likely seed desiccation responses, providing a gap-filling tool for global studies of plant traits, as well as critical decision-making support for plant conservation activities.

Maximizing the phylogenetic diversity of seed banks.

Ex situ conservation efforts such as those of zoos, botanical gardens, and seed banks will form a vital complement to in situ conservation actions over the coming decades. It is therefore necessary to pay the same attention to the biological diversity represented in ex situ conservation facilities as is often paid to protected-area networks. Building the phylogenetic diversity of ex situ collections will strengthen our capacity to respond to biodiversity loss. Since 2000, the Millennium Seed Bank Partnership has banked seed from 14% of the world's plant species. We assessed the taxonomic, geographic, and phylogenetic diversity of the Millennium Seed Bank collection of legumes (Leguminosae). We compared the collection with all known legume genera, their known geographic range (at country and regional levels), and a genus-level phylogeny of the legume family constructed for this study. Over half the phylogenetic diversity of legumes at the genus level was represented in the Millennium Seed Bank. However, pragmatic prioritization of species of economic importance and endangerment has led to the banking of a less-than-optimal phylogenetic diversity and prioritization of range-restricted species risks an underdispersed collection. The current state of the phylogenetic diversity of legumes in the Millennium Seed Bank could be substantially improved through the strategic banking of relatively few additional taxa. Our method draws on tools that are widely applied to in situ conservation planning, and it can be used to evaluate and improve the phylogenetic diversity of ex situ collections.

Drying banana seeds for ex situ conservation.

The ability of seeds to withstand drying is fundamental to ex situ seed conservation but drying responses are not well known for most wild species including crop wild relatives. We look at drying responses of seeds of Musa acuminata and Musa balbisiana, the two primary wild relatives of bananas and plantains, using the following four experimental approaches: (i) We equilibrated seeds to a range of relative humidity (RH) levels using non-saturated lithium chloride solutions and subsequently measured moisture content (MC) and viability. At each humidity level we tested viability using embryo rescue (ER), tetrazolium chloride staining and germination in an incubator. We found that seed viability was not reduced when seeds were dried to 4% equilibrium relative humidity (eRH; equating to 2.5% MC). (ii) We assessed viability of mature and less mature seeds using ER and germination in the soil and tested responses to drying. Findings showed that seeds must be fully mature to germinate and immature seeds had negligible viability. (iii) We dried seeds extracted from ripe/unripe fruit to 35-40% eRH at different rates and tested viability with germination tests in the soil. Seeds from unripe fruit lost viability when dried and especially when dried faster; seeds from ripe fruit only lost viability when fast dried. (iv) Finally, we dried and re-imbibed mature and less mature seeds and measured embryo shrinkage and volume change using X-ray computer tomography. Embryos of less mature seeds shrank significantly when dried to 15% eRH from 0.468 to 0.262 mm3, but embryos of mature seeds did not. Based on our results, mature seeds from ripe fruit are desiccation tolerant to moisture levels required for seed genebanking but embryos from immature seeds are mechanistically less able to withstand desiccation, especially when water potential gradients are high.

Corrigendum to: Drying banana seeds for ex situ conservation.

[This corrects the article DOI: 10.1093/conphys/coab099.].

Selecting the best candidates for resurrecting extinct-in-the-wild plants from herbaria.

Resurrecting extinct species is a fascinating and challenging idea for scientists and the general public. Whereas some theoretical progress has been made for animals, the resurrection of extinct plants (de-extinction sensu lato) is a relatively recently discussed topic. In this context, the term 'de-extinction' is used sensu lato to refer to the resurrection of 'extinct in the wild' species from seeds or tissues preserved in herbaria, as we acknowledge the current impossibility of knowing a priori whether a herbarium seed is alive and can germinate. In plants, this could be achieved by germinating or in vitro tissue-culturing old diaspores such as seeds or spores available in herbarium specimens. This paper reports the first list of plant de-extinction candidates based on the actual availability of seeds in herbarium specimens of globally extinct plants. We reviewed globally extinct seed plants using online resources and additional literature on national red lists, resulting in a list of 361 extinct taxa. We then proposed a method of prioritizing candidates for seed-plant de-extinction from diaspores found in herbarium specimens and complemented this with a phylogenetic approach to identify species that may maximize evolutionarily distinct features. Finally, combining data on seed storage behaviour and longevity, as well as specimen age in the novel 'best de-extinction candidate' score (DEXSCO), we identified 556 herbarium specimens belonging to 161 extinct species with available seeds. We expect that this list of de-extinction candidates and the novel approach to rank them will boost research efforts towards the first-ever plant de-extinction.

Seed Banks as Incidental Fungi Banks: Fungal Endophyte Diversity in Stored Seeds of Banana Wild Relatives.

Seed banks were first established to conserve crop genetic diversity, but seed banking has more recently been extended to wild plants, particularly crop wild relatives (CWRs) (e.g., by the Millennium Seed Bank (MSB), Royal Botanic Gardens Kew). CWRs have been recognised as potential reservoirs of beneficial traits for our domesticated crops, and with mounting evidence of the importance of the microbiome to organismal health, it follows that the microbial communities of wild relatives could also be a valuable resource for crop resilience to environmental and pathogenic threats. Endophytic fungi reside asymptomatically inside all plant tissues and have been found to confer advantages to their plant host. Preserving the natural microbial diversity of plants could therefore represent an important secondary conservation role of seed banks. At the same time, species that are reported as endophytes may also be latent pathogens. We explored the potential of the MSB as an incidental fungal endophyte bank by assessing diversity of fungi inside stored seeds. Using banana CWRs in the genus Musa as a case-study, we sequenced an extended ITS-LSU fragment in order to delimit operational taxonomic units (OTUs) and used a similarity and phylogenetics approach for classification. Fungi were successfully detected inside just under one third of the seeds, with a few genera accounting for most of the OTUs-primarily Lasiodiplodia, Fusarium, and Aspergillus-while a large variety of rare OTUs from across the Ascomycota were isolated only once. Fusarium species were notably abundant-of significance in light of Fusarium wilt, a disease threatening global banana crops-and so were targeted for additional sequencing with the marker EF1α in order to delimit species and place them in a phylogeny of the genus. Endophyte community composition, diversity and abundance was significantly different across habitats, and we explored the relationship between community differences and seed germination/viability. Our results show that there is a previously neglected invisible fungal dimension to seed banking that could well have implications for the seed collection and storage procedures, and that collections such as the MSB are indeed a novel source of potentially useful fungal strains.

Correlated evolution of genome size and seed mass.

Previous investigators have identified strong positive relationships between genome size and seed mass within species, and across species from the same genus and family. Here, we make the first broad-scale quantification of this relationship, using data for 1222 species, from 139 families and 48 orders. We analyzed the relationship between genome size and seed mass using a statistical framework that included four different tests. A quadratic relationship between genome size and seed mass appeared to be driven by the large genome/seed mass gymnosperms and the many small genome size/large seed mass angiosperms. Very small seeds were never associated with very large genomes, possibly indicating a developmental constraint. Independent contrast results showed that divergences in genome size were positively correlated with divergences in seed mass. Divergences in seed mass have been more closely correlated with divergences in genome size than with divergences in other morphological and ecological variables. Plant growth form is the only variable examined thus far that explains a greater proportion of variation in seed mass than does genome size.

A brief history of seed size.

Improved phylogenies and the accumulation of broad comparative data sets have opened the way for phylogenetic analyses to trace trait evolution in major groups of organisms. We arrayed seed mass data for 12,987 species on the seed plant phylogeny and show the history of seed size from the emergence of the angiosperms through to the present day. The largest single contributor to the present-day spread of seed mass was the divergence between angiosperms and gymnosperms, whereas the widest divergence was between Celastraceae and Parnassiaceae. Wide divergences in seed size were more often associated with divergences in growth form than with divergences in dispersal syndrome or latitude. Cross-species studies and evolutionary theory are consistent with this evidence that growth form and seed size evolve in a coordinated manner.

Factors that shape seed mass evolution.

We used correlated divergence analysis to determine which factors have been most closely associated with changes in seed mass during seed plant evolution. We found that divergences in seed mass have been more consistently associated with divergences in growth form than with divergences in any other variable. This finding is consistent with the strong relationship between seed mass and growth form across present-day species and with the available data from the paleobotanical literature. Divergences in seed mass have also been associated with divergences in latitude, net primary productivity, temperature, precipitation, and leaf area index. However, these environmental variables had much less explanatory power than did plant traits such as seed dispersal syndrome and plant growth form.