Surprisingly Weak Coordination between Leaf Structure and Function among Closely-related 1 Tomato Species 2 3

15 Natural selection may often favor coordination between different traits, or phenotypic 16 integration, in order to most efficiently acquire and deploy scarce resources. As leaves are the 17 primary photosynthetic organ in plants, many have proposed that leaf physiology, biochemistry, 18 and anatomical structure are coordinated along a functional trait spectrum from fast, resource-19 acquisitive syndromes to slow, resource-conservative syndromes. However, the coordination 20 hypothesis has rarely been tested at a phylogenetic scale most relevant for understanding rapid 21 adaptation in the recent past or predicting evolutionary trajectories in response to climate change. 22 To that end, we used a common garden to examine genetically-based coordination between leaf 23 traits across 19 wild and cultivated tomato taxa. We found surprisingly weak integration between 24 photosynthetic rate, leaf structure, biochemical capacity, and CO 2 diffusion, even though all were 25 arrayed in the predicted direction along a 'fast-slow' spectrum. This suggests considerable scope 26 for unique trait combinations to evolve in response to new environments or in crop breeding. In 27 particular, we find that partially independent variation in stomatal and mesophyll conductance 28 may allow a plant to improve water-use efficiency without necessarily sacrificing maximum 29 photosynthetic rates. Our study does not imply that functional trait spectra or tradeoffs are 30 unimportant, but that the many important axes of variation within a taxonomic group may be 31 unique and not generalizable to other taxa.