Interspecific morphological variation in Juglandoideae resting bud organization: a winter's tale?

BACKGROUND AND AIMS: Dormant resting buds are frequently regarded as static units, with protective cataphylls on the outside and embryonic foliage leaves on the inside. How the presence of cataphylls influences the dynamic, cyclical, annually repeating sequence of leaf forms that a resting bud gives rise to has rarely been interrogated. To examine the connection between dormant structure and growing-season development, we compare the complete seasonal heteroblastic sequence of leaf forms of six species of temperate Juglandaceae with distinctly different vegetative resting bud structures. These include buds with cataphylls; buds without cataphylls; and buds with caducous cataphylls that are lost before the onset of winter. METHODS: In a common garden setting over a 7-month growing season, the dimensions of 2249 individual vegetative metamers were tracked from first exposure to abscission along the shoots of saplings and mature trees. The timing of metamer initiation within terminal buds was investigated using micro-CT scanning. Character state transitions of resting bud types were estimated using a phylogenetic tree of Juglandaceae. KEY RESULTS: The presence of cataphylls within a heteroblastic sequence is associated with a single cohort of foliage leaves that flush and abscise synchronously. This growing pattern is highly determinate, with next year's terminal-bud cataphylls already initiated before spring leaf out. In contrast, in sequences without cataphylls, shorter-lived foliage leaves appear and abscise in a staggered fashion. Despite these differences in leaf demography, all examined heteroblastic sequences produce a series of small, caducous leaf forms that precede terminal bud set. CONCLUSIONS: The ubiquity of caducous leaf forms in Juglandoideae may point to the importance of shoot tip protection far beyond the dormant season. In addition, the presence or absence of cataphylls in resting buds is indicative of distinct shoot ontogenetic patterns, and functional strategies, in summer.

Naked resting bud morphologies and their taxonomic and geographic distributions in temperate, woody floras.

Resting bud cataphylls are often assumed to play an essential protective role in winter due to their widespread presence among temperate, woody plants. This view is challenged by our documentation of significant numbers of temperate woody angiosperm taxa with naked buds that overwinter without cataphyll protection. We inventoried temperate, woody angiosperm taxa reported to have resting buds without cataphyll protection in winter and for the first time characterised the morphological and functional diversity of naked buds. Using this new classification of bud types, the taxonomic and geographic distributions of taxa with naked buds were summarised and relationships between plant functional traits and bud type were investigated. Naked buds are not, as long presumed, markedly rare in temperate, woody floras. They occur in at least 87 genera in 42 families throughout the angiosperm phylogeny in various morphologically distinct manifestations. The geographic distribution of species with naked buds in temperate areas was found to be associated with summer precipitation, but not with winter climatic variables. Resting bud structure is not necessarily a trait optimised solely for winter survival. A taxon's bud composition may be influenced by factors such as biogeographic history and ontogenetic pattern of leaf formation over the growing season.

The ancestral flower of angiosperms and its early diversification.

Recent advances in molecular phylogenetics and a series of important palaeobotanical discoveries have revolutionized our understanding of angiosperm diversification. Yet, the origin and early evolution of their most characteristic feature, the flower, remains poorly understood. In particular, the structure of the ancestral flower of all living angiosperms is still uncertain. Here we report model-based reconstructions for ancestral flowers at the deepest nodes in the phylogeny of angiosperms, using the largest data set of floral traits ever assembled. We reconstruct the ancestral angiosperm flower as bisexual and radially symmetric, with more than two whorls of three separate perianth organs each (undifferentiated tepals), more than two whorls of three separate stamens each, and more than five spirally arranged separate carpels. Although uncertainty remains for some of the characters, our reconstruction allows us to propose a new plausible scenario for the early diversification of flowers, leading to new testable hypotheses for future research on angiosperms.