Pre-Carpels from the Middle Triassic of Spain

In stark contrast to the multitude of hypotheses on carpel evolution, there is little fossil evidence testing these hypotheses. The recent discovery of angiosperms from the Early Jurassic makes the search for precursors of angiosperm carpels in the Triassic more promising. Our light microscopic and SEM observations on Combina gen. nov., a cone-like organ from the Middle Triassic of Spain, indicate that its lateral unit includes an axillary anatropous ovule and a subtending bract, and the latter almost fully encloses the former. Such an observation not only favors one of the theoretical predictions but also makes some Mesozoic gymnosperms (especially conifers and Combina) comparable to some angiosperms. Combina gen. nov. appears to be an important chimeric fossil plant that may complete the evidence chain of the origin of carpels in geological history, partially narrowing the gap between angiosperms and gymnosperms.