Pre-Gondwanan-breakup origin of Beauprea (Proteaceae) explains its historical presence in New Caledonia and New Zealand

New Caledonia and New Zealand belong to the now largely submerged continent Zealandia. Their high levels of endemism and species richness are usually considered the result of transoceanic dispersal events followed by diversification after they re-emerged from the Pacific Ocean in the mid-Cenozoic. We explore the origin and evolutionary history of Beauprea (Proteaceae), which is now endemic to New Caledonia but was once spread throughout eastern Gondwana, including New Zealand. We review the extensive Beauprea-type pollen data in the fossil records and analyze the relationship of these fossil taxa to extant genera within Proteaceae. We further reconstruct the phylogenetic relations among nine extant species of Beauprea and estimate the age of the Beauprea clade. By incorporating extinct taxa into the Beauprea phylogenetic tree, we reconstruct the ancient distribution of this genus. Our analysis shows that Beauprea originated c. 88 Ma (million years ago) in Antarctica–Southeastern Australia and spread throughout Gondwana before its complete breakup. We propose that Beauprea, already existing as two lineages, was carried with Zealandia when it separated from the rest of Gondwana c. 82 Ma, thus supporting an autochthonous origin for Beauprea species now in New Caledonia and historically in New Zealand up to 1 Ma. We show that the presence of Beauprea through transoceanic dispersal is implausible. This means that neither New Caledonia nor New Zealand has been entirely submerged since the Upper Cretaceous; thus, possible vicariance and allopatry must be taken into account when considering the high levels of endemism and species richness of these island groups

A Cretaceous origin for fire adaptations in the Cape flora

Fire has had a profound effect on the evolution of worldwide biotas. The Cape Floristic Region is one of the world’s most species-rich regions, yet it is highly prone to recurrent fires and fire-adapted species contribute strongly to the overall flora. It is hypothesized that the current fire regimes in the Cape could be as old as 6–8 million years (My), while indirect evidence indicates that the onset of fire could have reached 18 million years ago (Ma). Here, we trace the origin of fire-dependent traits in two monocot families that are significant elements in the fire-prone Cape flora. Our analysis shows that fire-stimulated flowering originated in the Cape Haemodoraceae 81 Ma, while fire-stimulated germination arose in the African Restionaceae at least 70 Ma, implying that wildfires have been a significant force in the evolution of the Cape flora at least 60 My earlier than previous estimates. Our results provide strong evidence for the presence of fire adaptations in the Cape from the Cretaceous, leading to the extraordinary persistence of a fire-adapted flora in this biodiversity hotspot, and giving support to the hypothesis that Cretaceous fire was a global phenomenon that shaped the evolution of terrestrial floras

The role of pollinators in the floral diversification and life history of Leucospermum (Proteaceae)

Leucospermum, commonly known as pincushions, are an endemic genus often associated with South Africa and, more specifically, the fynbos ecosystem and Cape Floral Kingdom. The staggering floral polymorphism within the genus suggests a wide variety of pollination modes. Despite this remarkable assortment of floral morphologies within Leucospermum only recently has any research been conducted to explain the role of pollinators in driving diversification. In this study we construct the first molecular phylogeny for the genus and, taking relatedness into account using a phylogenetic generalized least squares approach, determine how pollinators have driven floral divergence. Pollinators were assigned through direct observations and a suite of floral traits that are likely to reflect pollination mode were measured for each taxa. In addition, the ability to autonomously self-pollinate was determined for each taxon and used to test if autogamy is linked to specific classes of pollinators, specialized taxa or recently derived taxa. All fieldwork has been finalized and we are awaiting our phylogenetic analysis before any results can be determined

Seed size, fecundity and postfire regeneration strategy are interdependent in Hakea

Seed size is a key functional trait that affects plant fitness at the seedling stage and may vary greatly with species fruit size, growth form and fecundity. Using structural equation modelling (SEM) and correlated trait evolution analysis, we investigated the interaction network between seed size and fecundity, postfire regeneration strategy, fruit size, plant height and serotiny (on-plant seed storage) among 82 species of the woody shrub genus, Hakea, with a wide spectrum of seed sizes (2–500 mg). Seed size is negatively correlated with fecundity, while fire-killed species (nonsprouters) produce more seeds than resprouters though they are of similar size. Seed size is unrelated to plant height and level of serotiny while it scales allometrically with fruit size. A strong phylogenetic signal in seed size revealed phylogenetic constraints on seed size variation in Hakea. Our analyses suggest a causal relationship between seed size, fecundity and postfire regeneration strategy in Hakea. These results demonstrate that fruit size, fecundity and evolutionary history have had most control over seed size variation among Hakea species

Small-seeded Hakea species tolerate cotyledon loss better than large-seeded congeners

Six Hakea species varying greatly in seed size were selected for cotyledon damage experiments. The growth of seedlings with cotyledons partially or completely removed was monitored over 90 days. All seedlings perished by the fifth week when both cotyledons were removed irrespective of seed size. Partial removal of cotyledons caused a significant delay in the emergence of the first leaf, and reduction in root and shoot growth of the large-seeded species. The growth of seedlings of small-seeded species was less impacted by cotyledon damage. The rate of survival, root and shoot lengths and dry biomass of the seedlings were determined after 90 days. When seedlings were treated with balanced nutrient solutions following removal of the cotyledons, survival was 95–98%, but 0% when supplied with nutrient solutions lacking N or P or with water only. The addition of a balanced nutrient solution failed to restore complete growth of any species, but the rate of root elongation for the small-seeded species was maintained. Cotyledons provide nutrients to support early growth of Hakea seedlings, but other physiological roles for the cotyledons are also implicated. In conclusion, small-seeded Hakea species can tolerate cotyledons loss better than large-seeded species

Fire-mediated germination syndromes in Leucadendron (Proteaceae) and their functional correlates

A mechanistic understanding of fire-driven seedling recruitment is essential for effective conservation management of fire-prone vegetation, such as South African fynbos, especially with rare and threatened taxa. The genus Leucadendron (Proteaceae) is an ideal candidate for comparative germination studies, comprising 85 species with a mixture of contrasting life-history traits (killed by fire vs able to resprout; serotinous vs geosporous) and seed morphologies (nutlets vs winged achenes). Individual and combined effects of heat and smoke on seed germination of 40 species were quantified in the laboratory, and Bayesian inference applied to distinguish biologically meaningful treatment effects from non-zero, but biologically trivial, effects. Three germination syndromes were identified based on whether germination was dependent on, enhanced by, or independent of direct fire cues (heat and smoke). Seed storage location was the most reliable predictor of germination syndromes, with soil-stored seeds c. 80% more likely to respond to direct fire cues (primarily smoke) than canopy-stored seeds. Notable exceptions were L. linifolium, with an absolute requirement for smoke to germinate (the third serotinous species so reported), and two other serotinous species with smoke-enhanced germination. Nutlet-bearing species, whether serotinous or geosporous, were c. 70% more likely to respond to fire cues than winged seeds, but there was no evidence for an effect of phylogeny or persistence strategy on germination. This comprehensive account of seed germination characteristics and identification of germination syndromes and their predictors, supports propagation, conservation and restoration initiatives in this iconic fynbos genus and other fire-prone shrubs with canopy or soil-stored seeds

Biological and geophysical feedbacks with fire in the Earth system

Roughly 3% of the Earth's land surface burns annually, representing a critical exchange of energy and matter between the land and atmosphere via combustion. Fires range from slow smouldering peat fires, to low-intensity surface fires, to intense crown fires, depending on vegetation structure, fuel moisture, prevailing climate, and weather conditions. While the links between biogeochemistry, climate and fire are widely studied within Earth system science, these relationships are also mediated by fuels—namely plants and their litter—that are the product of evolutionary and ecological processes. Fire is a powerful selective force and, over their evolutionary history, plants have evolved traits that both tolerate and promote fire numerous times and across diverse clades. Here we outline a conceptual framework of how plant traits determine the flammability of ecosystems and interact with climate and weather to influence fire regimes. We explore how these evolutionary and ecological processes scale to impact biogeochemical and Earth system processes. Finally, we outline several research challenges that, when resolved, will improve our understanding of the role of plant evolution in mediating the fire feedbacks driving Earth system processes. Understanding current patterns of fire and vegetation, as well as patterns of fire over geological time, requires research that incorporates evolutionary biology, ecology, biogeography, and the biogeosciences

Post-fire litter microsites: Safe for seeds, unsafe for seedlings

We explore the effect of post—fire microsites on seed and seedling distribution and hence their potential role in community restoration. A summer wildfire and control burn in a sclerophyll shrubland in mediterranean Australia produced mosaics of physically and chemically contrasting microsites of litter and sand. Most seeds (>75%) of all species released from the burnt canopies fell, or were redispersed by wind, into the litter patches after both fires. Data on microsite characteristics and wind exposure (fire intensity), height of fruits, time of release, and seed properties were required to interpret relative distribution between the litter and sand patches. Seeds remained equally viable (up to 100%) over summer—autumn in the litter and sand and had equally high rates and levels (up to 100%) of subsequent winter germination. However, seedlings were 2—3 times less likely to survive in the litter and survivors were 35% smaller than those in the sand by the end of the first summer. Banksia hookeriana was particularly vulnerable to microsite properties, whereas the needle—leaved Hakea polyathema showed only minor responses. Pre—summer thinning of seedlings in the litter increased survival of the remainder by 2 times and size of the survivors by 31%. The fire—sensitive, small—seeded B. hookeriana had 17 times more seeds in the backburn litter than the resprouting, larger—seeded B. attenuata, which more than compensated for its 3 times greater seedling mortality levels over the dry summer. Recruitment of species prone to density—dependent mortality in the litter was enhanced by the retention of some seeds in the sand where competition for water was minimal, as indicated by the 2.2 times greater stomal conductance of their seedlings in early summer

Grasstree stem analysis reveals insufficient data for inference of fire history

Grinding back dead leaf bases on the stems of arborescent grasstrees (Xant7wrrlwea spp.) reveals a pattern of horizontal bands that has been interpreted as a record of the fire history experienced by the plant. The validity of this fire history record has previously been assessed through comparison of 100 grasstree stems sampled from shrubland near Eneabba in Western Australia against a 30 year fire history determined from satellite imagery. This analysis showed that the two records matched more than would be expected by chance, but concluded that the interpretation of the grasstree record as a fire history was not warranted as most of the grasstree fire records did not match satellite fire records. A second analysis of the same two sets of records, published in this journal, also showed that the records matched more than by chance, but concluded that the interpretation of grasstree banding as fire history was valid, though it failed to quantify the strength of this agreement. Here we examine' the approaches and interpretations of the two previously published studies, and provide new analyses to refine estimates of the amount of fire-related data present in the grasstree record. We show that only ~20% of grasstree 'fire' records may be attributable to fire. With eight out of ten of records not attributable to fire, we confirm that the grasstree record in its current form cannot be interpreted as fire history, and therefore claims of the grasstree technique to support management actions are untenable