19We reconsider the existing paradigm and show that the symbiotic options available to 20 the first plants emerging onto the land were more varied than previously thought.
22The current paradigm 23 The colonisation of the terrestrial environment by plants was a major turning point in Earth's carbohydrates fixed from atmospheric carbon dioxide through photosynthesis [5].
35Plant life diversified on land some 70 MY after the diversification of most major 36 animal lineages in the seas during the Cambrian explosion ( Fig. 1) [6]. Classic palaeoclimate 37 3 modelling shows that land plants diversified against a backdrop of falling atmospheric CO 2
38[7] (Figure 2), likely driven by growing demand and evolving capacity for carbon 39 assimilation of the burgeoning Earth flora [8,9]. Long before land plants emerged, however, 40 the terrestrial environment had been colonised by fungi [10,11]. Among the early-branching 41 fungal lineages were those that today form mutualistic associations with most plants (Figure 42 1). The macrofossil record for non-vascular plants is even more fragmentary than that for 43 vascular plants, though the study of microfossils is now providing key new data [12,13] lineages of land plants [24,25].
60The resulting paradigm is that the earliest, rootless, terrestrial plants co-evolved with
61Glomeromycota fungi [4,[26][27][28][29][30]
153We now know that liverwort-Glomeromycota symbioses can be mutualistic, involving fungi within plant tissues. An alternative but untested hypothesis is that their evolution was 287 linked to harbouring an overlooked or extinct microbial symbiont of green algae [22].
288The assumption that function of mycorrhizal and mycorrhiza-like fungal associations 289 is also conserved across land plant evolution has, until recently, been based on striking However, while we know for instance that functioning in modern vascular plant-