Insular woodiness refers to the evolutionary transition from herbaceousness to- wards derived woodiness on (sub)tropical islands and leads to island floras that have a higher proportion of woody... Show moreInsular woodiness refers to the evolutionary transition from herbaceousness to- wards derived woodiness on (sub)tropical islands and leads to island floras that have a higher proportion of woody species compared to floras of nearby continents.Several hypotheses have tried to explain insular woodiness since Darwin’s original observations, but experimental evidence why plants became woody on islands is scarce at best.Here, we combine experimental measurements of hydraulic failure in stems (as a proxy for drought stress resistance) with stem anatomical observations in the daisy lineage (Asteraceae), including insular woody Argyranthemum species from the Canary Islands and their herbaceous continental relatives.Our results show that stems of insular woody daisies are more resistant to drought- induced hydraulic failure than the stems of their herbaceous counterparts. The ana- tomical character that best predicts variation in embolism resistance is intervessel pit membrane thickness (TPM), which can be functionally linked with air bubble dynamics throughout the 3D vessel network. There is also a strong link between TPM vs. degree of woodiness and thickness of the xylem fibre wall vs. embolism resistance, resulting in an indirect link between lignification and resistance to embolism formation.Thicker intervessel pit membranes in Argyranthemum functionally explain why this insular woody genus is more embolism resistant to drought-induced failure compared to the herbaceous relatives from which it has evolved, but additional data are needed to confirm that palaeoclimatic drought conditions have triggered wood formation in this daisy lineage. Show less
The water transport pipeline in herbs is assumed to be more vulnerable to drought than in trees due to the formation of frequent embolisms (gas bubbles), which could be removed by the... Show moreThe water transport pipeline in herbs is assumed to be more vulnerable to drought than in trees due to the formation of frequent embolisms (gas bubbles), which could be removed by the occurrence of root pressure, especially in grasses. Here, we studied hydraulic failure in herbaceous angiosperms by measuring the pressure inducing 50% loss of hydraulic conductance (P50) in stems of 26 species, mainly European grasses (Poaceae). Our measurements show a large range in P50 from 20.5 to 27.5 MPa, which overlaps with 94% of the woody angiosperm species in a worldwide, published data set and which strongly correlates with an aridity index. Moreover, the P50 values obtained were substantially more negative than the midday water potentials for five grass species monitored throughout the entire growing season, suggesting that embolism formation and repair are not routine and mainly occur under water deficits. These results show that both herbs and trees share the ability to withstand very negative water potentials without considerable embolism formation in their xylem conduits during drought stress. In addition, structure-function trade-offs in grass stems reveal that more resistant species are more lignified, which was confirmed for herbaceous and closely related woody species of the daisy group (Asteraceae). Our findings could imply that herbs with more lignified stems will become more abundant in future grasslands under more frequent and severe droughts, potentially resulting in lower forage digestibility. Show less
The water transport pipeline in herbs is assumed to be more vulnerable to drought than in trees due to the formation of frequent embolisms (gas bubbles), which could be removed by the occurrence of... Show moreThe water transport pipeline in herbs is assumed to be more vulnerable to drought than in trees due to the formation of frequent embolisms (gas bubbles), which could be removed by the occurrence of root pressure, especially in grasses. Here, we studied hydraulic failure in herbaceous angiosperms by measuring the pressure inducing 50% loss of hydraulic conductance (P50)in stems of 26 species, mainly European grasses (Poaceae). Our measurements show a large range in P50 from 20.5 to 27.5 MPa, which overlaps with 94% of the woody angiosperm species in a worldwide, published data set and which strongly correlates with an aridity index. Moreover, the P50 values obtained were substantially more negative than the midday water potentials for five grass species monitored throughout the entire growing season, suggesting that embolism formation and repair are not routine and mainly occur under water deficits. These results show that both herbs and trees share the ability to withstand very negative water potentials without considerable embolism formation in their xylem conduits during drought stress. In addition, structure-function trade-offs in grass stems reveal that more resistant species are more lignified, which was confirmed for herbaceous and closely related woody species of the daisy group (Asteraceae). Our findings could imply that herbs with more lignified stems will become more abundant in future grasslands under more frequent and severe droughts, potentially resulting in lower forage digestibility. Show less
Lens, F.P.; Davin, N.F.; Smets, E.F.; Arco, M. del 2013
Premise of research. One of the most conspicuous aspects of island floras is the relatively high proportion of woody species. Often, but not always, these woody species have developed wood on the... Show morePremise of research. One of the most conspicuous aspects of island floras is the relatively high proportion of woody species. Often, but not always, these woody species have developed wood on the islands and have evolved from herbaceous continental ancestors, a phenomenon known as insular woodiness. Shifts from herbaceousness toward increased woodiness also occur on continents (the broader term “secondary woodiness” is more appropriate here and includes insular woodiness), but comprehensive worldwide knowledge about secondary woodiness within angiosperms remains lacking. We update hypotheses regarding the herbaceous ancestry of woody Canarian lineages in a molecular phylogenetic context and investigate the possible link of secondary woodiness and paedomorphic wood features in the Carlquistian sense. Methodology. We have assembled available literature data from molecular phylogenetic studies, wood anatomical descriptions, floras, and taxonomic revisions to identify the native secondarily woody taxa. Pivotal results. In total, at least 220 native Canary Island species of flowering plants, from 34 genera representing 15 families, are truly insular woody. This represents a significant portion of the native nonmonocot angiosperm species on the Canaries, and all of the insular woody species have paedomorphic wood features in the Carlquistian sense, although this wood anatomical syndrome might be more related to particular life forms. The majority of these insular woody groups typically grow in the markedly dry lowland regions, suggesting a possible link between secondary woodiness and increased drought resistance. Conclusions. The Canary Island flora is characterized by at least 38 independent shifts toward insular woodiness, representing an important portion of the endemic angiosperms on the archipelago. These convergent evolutionary events emphasize the remarkable lability in growth forms between herbaceous and woody lineages, but it remains puzzling which environmental variables trigger these shifts and how these independent shifts are regulated genetically. Show less
