RESUMEN
Frost tolerance has evolved many times independently across flowering plants. However, conservation of several frost tolerance mechanisms among distant relatives suggests that apparently independent entries into freezing climates may have been facilitated by repeated modification of existing traits ('precursor traits'). One possible precursor trait for freezing tolerance is drought tolerance, because palaeoclimatic data suggest plants were exposed to drought before frost and several studies have demonstrated shared physiological and genetic responses to drought and frost stress. Here, we combine ecophysiological experiments and comparative analyses to test the hypothesis that drought tolerance acted as a precursor to frost tolerance in cool-season grasses (Pooideae). Contrary to our predictions, we measured the highest levels of frost tolerance in species with the lowest ancestral drought tolerance, suggesting that the two stress responses evolved independently in different lineages. We further show that drought tolerance is more evolutionarily labile than frost tolerance. This could limit our ability to reconstruct the order in which drought and frost responses evolved relative to each other. Further research is needed to determine whether our results are unique to Pooideae or general for flowering plants.
RESUMEN
BACKGROUND: Variation in floral shapes has long fascinated biologists and its modelling enables testing of evolutionary hypotheses. Recent comparative studies that explore floral shape have largely ignored 3D floral shape. We propose quantifying floral shape by using geometric morphometrics on a virtual3D model reconstructed from 2D photographical data and demonstrate its performance in capturing shape variation. METHODS: This approach offers unique benefits to complement established imaging techniques (i) by enabling adequate coverage of the potential morphospace of large and diverse flowering-plant clades; (ii) by circumventing asynchronicity in anthesis of different floral parts; and (iii) by incorporating variation in copy number of floral organs within structures. We demonstrate our approach by analysing 90 florally-diverse species of the Southern African genus Pelargonium (Geraniaceae). We quantify Pelargonium floral shapes using 117 landmarks and show similarities in reconstructed morphospaces for nectar tube, corolla (2D datasets), and a combined virtual3D dataset. RESULTS: Our results indicate that Pelargonium species differ in floral shape, which can also vary extensively within a species. PCA results of the reconstructed virtual3D floral models are highly congruent with the separate 2D morphospaces, indicating it is an accurate, virtual, representation of floral shape. Through our approach, we find that adding the third dimension to the data is crucial to accurately interpret the manner of, as well as levels of, shape variation in flowers.