Both engineered hydraulic systems and plant hydraulic systems are protected against failure by resistance, reparability, and redundancy. A basic rule of reliability engineering is that the level of independent redundancy should increase with increasing risk of fatal system failure. Here we show that hydraulic systems of plants function as predicted by this engineering rule. Hydraulic systems of shrubs sampled along two transcontinental aridity gradients changed with increasing aridity from highly integrated to independently redundant modular designs. Shrubs in humid environments tend to be hydraulically integrated, with single, round basal stems, whereas dryland shrubs typically have modular hydraulic systems and multiple, segmented basal stems. Modularity is achieved anatomically at the vessel-network scale or developmentally at the whole-plant scale through asymmetric secondary growth, which results in a semiclonal or clonal shrub growth form that appears to be ubiquitous in global deserts.plant hydraulic systems ͉ wood anatomy ͉ hydraulic redundancy ͉ xylem structure and function I n engineering terms, the hydraulic system of a plant is a negative-pressure flow system. This type of hydraulic system, whether natural or man-made, is prone to fail when air bubbles (emboli) are introduced, because under strong negative pressure a single embolism can lead to breakage of the water column unless the air bubble is isolated in a branch or pipe. Both drought and freezing can cause embolisms in plants (1).Drought-induced embolisms form under negative pressure, when air is pulled into a water-filled conduit from adjacent air-filled spaces or cells, a process known as ''air seeding.'' This common, even daily, event (2-4) can lead to complete failure of the hydraulic system if runaway embolism occurs (5). Two of the three attributes by which plants' negative-pressure flow systems can be protected against failure, resistance and reparability, have been subjects of active research during the last decade (2-4, 6-10). The third attribute, redundancy, has received much less attention as an important drought adaptation but is emerging as a focus of research (11)(12)(13)(14). Attributes of redundancy in hydraulic systems of vessel-bearing angiosperms include the numbers of vessels (14), the vessel network topology (12), the number and sizes of pits between adjacent vessels (13,15,16), and the division of whole plants into independent hydraulic units (17).A basic rule of reliability engineering states that the level of independent redundancy should increase with increasing risk of fatal system failure (18); hydraulic engineers routinely increase the safety of man-made pressure-flow systems by designing them to be redundant (19). Redundancy in hydraulic systems (Fig. 1) can vary from a high degree of inter-connectedness (i.e., integrated redundancy) to complete, independent compartmentation (i.e., modular redundancy). In a negative-pressure flow system, integrated redundancy allows alternate water transport pathways around blockage...