19Natural selection is a major driver for the origins of adaptations and new species 1 . Whether or 20 not the processes driving adaptation and speciation share a molecular basis remains largely 21 unknown 2 . Here, we show that divergence in hormone signalling contributed to the evolution 22 habits of S. lautus populations, therefore making evolution of the auxin pathway a natural 60 candidate to link the molecular basis of adaptation and speciation. We reasoned that if 61 divergence in the auxin pathway contributed to the evolution of adaptation and speciation in 62 S. lautus, we would discover the following evidence: First, we would detect similar patterns 63 of genetic divergence in auxin-related pathways across multiple erect and prostrate hybrid 64 and natural populations. Second, these populations would differ in phenotypes dependent on 65 auxin, such as their ability to alter the direction of growth in relation to gravity 10,16 . And third, 66 divergence in these auxin-dependent phenotypes would contribute to local adaptation and 67 intrinsic reproductive isolation between populations. 68We test these hypotheses primarily on coastal populations of S. lautus (Fig. 1a, Extended 69Data Table 1), which exhibit strong correlations between growth habit and the environments 70 they occupy 7 . Populations inhabiting sand dunes (Dune hereafter) are erect, while populations 71 growing on adjacent rocky headlands (Headland hereafter) are prostrate ( Fig. 1b). Erect and 72 prostrate growth habits can also be found in related populations from the alpine regions of 73 Australia, with a prostrate population inhabiting an exposed alpine meadow and an erect 74 population inhabiting a sheltered alpine gully (Fig. 1c). Dune populations are continually 75 exposed to high temperatures and sun radiation, low salinity, and low nutrient sand substrate, 76whereas Headland populations are exposed to high salinity, high nutrients and powerful 77 winds 17 . Neighbouring Dune and Headland populations are often sister taxa, group into two 78 major monophyletic clades (eastern and south-eastern) and have evolved their contrasting 79 growth habits independently multiple times 7,20 . These Dune and Headland populations are 80 locally adapted [17][18][19][20] and their F2 hybrids have low fitness 21 , indicating the presence of 81 intrinsic reproductive isolation. Furthermore, performing genetic, physiological, and 82 ecological experimental studies is achievable in this system due to its short life cycle, diploid 83 inheritance, and small vegetative size. Therefore, the Senecio lautus species complex 84The physiological basis of repeated evolution in S. lautus 126Considering we identified a multitude of different auxin related genes between erect and 127 prostrate populations of S. lautus and the regulation and transport of auxin is well established 128 to modulate gravitropism in plants, we predicted that these divergent growth habits may be a 129 direct consequence of changes in the auxin pathway, and can therefore contribute to 130 d...