Given that stomatal movement is ultimately a mechanical process and that stomata are morphologically and mechanically diverse, we explored the influence of stomatal mechanical diversity on leaf gas exchange and considered some of the constraints. Mechanical measurements were conducted on the guard cells of four different species exhibiting different stomatal morphologies, including three variants on the classical ''kidney'' form and one ''dumb-bell'' type; this information, together with gas-exchange measurements, was used to model and compare their respective operational characteristics. Based on evidence from scanning electron microscope images of cryo-sectioned leaves that were sampled under full sun and high humidity and from pressure probe measurements of the stomatal aperture versus guard cell turgor relationship at maximum and zero epidermal turgor, it was concluded that maximum stomatal apertures (and maximum leaf diffusive conductance) could not be obtained in at least one of the species (the grass Triticum aestivum) without a substantial reduction in subsidiary cell osmotic (and hence turgor) pressure during stomatal opening to overcome the large mechanical advantage of subsidiary cells. A mechanism for this is proposed, with a corollary being greatly accelerated stomatal opening and closure. Gas-exchange measurements on T. aestivum revealed the capability of very rapid stomatal movements, which may be explained by the unique morphology and mechanics of its dumb-bell-shaped stomata coupled with ''see-sawing'' of osmotic and turgor pressure between guard and subsidiary cells during stomatal opening or closure. Such properties might underlie the success of grasses.Although the morphological diversity of stomata is widely documented (Haberlandt, 1884;Meidner and Mansfield, 1968;Allaway and Milthorpe, 1976;Ziegler, 1987;Willmer and Fricker, 1996), little is known of how this translates into functional diversity and what the environmental context of this might be. Throughout the 400 million year (Ma) history of vascular plants on land, long-term decline in atmospheric CO 2 concentration and shifts in prevailing moisture patterns have placed selective pressures on stomata to increase epidermal conductance to CO 2 diffusion and also to increase transpiration efficiency (CO 2 fixed per unit water transpired). This posed two separate problems, upon which the combination of mutation and time might have worked to give rise to the current diversity of stomatal form and function. The first centered on the simple geometric practicalities of fitting enough functional stomatal units per unit leaf surface area to meet the desired CO 2 flux as atmospheric CO 2 concentration changed, or to service an increase in photosynthetic capacity. The second centered on the performance characteristics of any new stomatal structure or configuration in relation to transpiration efficiency. Here, by examining the mechanical and performance characteristics of stomata in four different species, we explore the nature of these two problem...