Exploring the causes of variation in phenology and morphology in Mediterranean geophytes: a genus-wide study of Cyclamen

The phenology and morphology of Mediterranean plants are constrained by drought in summer and cold temperatures in winter. In this study we examine how climatic factors and phylogenetic constraints have shaped variation in the phenology and morphology of 17 species of the genus Cyclamen cultivated in uniform garden conditions. We quantify the extent to which traits differ among subgenera and thus represent conserved traits within evolutionary lineages. We also explore whether leaf, flowering and seed-release phenology are correlated among species, and thus whether variation in flowering phenology results from selection on dispersal phenology. Our results show a significant influence of subgenus membership on leaf and flowering phenology but not on morphological traits or the timing of seed release. Among-species variation in foliage height, leaf size and seed mass (but not in floral traits) is correlated with chromosome number. Leaf traits show that species with a shorter vegetative period have a higher capacity for resource acquisition. Major phenological shifts, i.e. spring vs. autumn flowering and a decoupling of leaf and flower phenology in autumnal flowering species, thus occurred prior to the diversification of species in each subgenus and not as a response to selection on dispersal timing. Leaf and flowering phenology illustrate a gradient of strategies from autumn flowering in the absence of leaves (hysteranthous species) to spring flowering with fully developed foliage (synanthous species). In the former, flowering is uncoupled from resource acquisition by simultaneous photosynthesis, indicative that hysteranthy is a response to temporal unpredictability in the onset of rain after the summer drought. Our results support the idea that whereas leaf development is controlled primarily by moisture availability and secondarily by temperature, flowering is temperature dependent, above a minimum moisture threshold. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 145 , 469–484.     

Decoupling of reproduction and growth: an unusual pattern in the life cycle of the Mediterranean geophyte Narcissus serotinus

The unpredictable seasonality of the Mediterranean climate imposes on the development of specific life‐cycle phenologies such as those seen in geophyte species that have a period of dormancy to avoid stressful abiotic conditions. To date, two life‐cycle strategies are recognized in Mediterranean geophytes. In some species, growth and reproduction occur in different seasons (hysteranthous geophytes), whereas in others these two processes occur at the same time (synanthous geophytes). A study of the life cycle of the autumnal‐flowering Narcissus serotinus in three populations over two consecutive years confirmed that neither the hysteranthy nor the synanthy strategy could be applied to this geophyte. Rather, reproduction and growth occur simultaneously, but in different plants. A strong correlation was found between bulb weight and the presence of reproductive or vegetative plants.