This paper describes the 'recruitment box,' an integrative model that defines the stream stage patterns that enable successful establishment of riparian cottonwood seedlings. In western North America, cottonwood seed dispersal generally occurs after annual peak river flows. The receding stream exposes moist sites upon which seeds land after transport by wind and water, Germination is rapid, and initial seedling establishment is often prolific. However, the vast majority of seedlings die, primarily due to drought stress, as root growth is insufficient to maintain contact with the receding zone of moisture. Cottonwood roots grow about 0.5 to 1 cm per day or 60 to 100 cm in the first year. Along the 'losing' streams in semi-arid regions, the riparian water table is an almost horizontal extension from the stream stage. A capillary fringe exists above the water table and is often 30 to 40 cm in elevation, but can range from about 5 to 130 em depending on substrate texture. The combination of root growth and capillary fringe define the successful recruitment band, which is usually from about 0.6 to 2 m in elevation above the late summer stream stage. Within this range, higher elevation establishment occurs (i) for the Aigeiros cottonwoods, Populus dettoides, and P. fremontii, which grow more rapidly than Tacamahaca species and occur in warmer areas with longer growing seasons; (ii) along larger rivers that are characterized by more gradual stage fluctuations; and (iii) along streams with finer substrate. The rate of stream stage decline is also critical for seedling survival and should not exceed 2.5 cm per day. The recruitment box model is consistent with dendrochronological interpretations that moderate flood events are naturally required for cottonwood recruitment. Flood events with recurrences of about 1 in 5 to 1 in 10 years often satisfy the model and provide stream stage patterns with a gradual decline through the recruitment box. The model will facilitate analyses of the reproductive ecology of riparian cottonwoods and also permit the prescription of stream stage patterns for cottonwood seedling recruitment along danuned rivers.
Cottonwoods (Populus spp.) are adapted to riparian or floodplain zones throughout the Northern Hemisphere; they are also used as parents for fast-growing hybrid poplars. We review recent ecophysiological studies of the native cottonwoods Populus angustifolia James, P. balsamifera L., P. deltoides Marsh., P. fremontii S. Watson and P. trichocarpa T. & G. in North America, and P. nigra L. in Europe. Variation exists within and across species and hybrids; however, all riparian cottonwoods are dependent on shallow alluvial groundwater that is linked to stream water, particularly in semi-arid regions. This conclusion is based on studies of their natural occurrence, decline following river damming and dewatering (water removal), water relations, isotopic composition of xylem water, and by the establishment of cottonwoods along formerly barren natural channels after flow augmentation in response to the conveyance of irrigation water. When alluvial groundwater is depleted as a result of river dewatering or groundwater pumping, riparian cottonwoods exhibit drought-stress responses including stomatal closure and reduced transpiration and photosynthesis, altered 13C composition, reduced predawn and midday water potentials, and xylem cavitation. These physiological responses are accompanied by morphological responses including reduced shoot growth, altered root growth, branch sacrifice and crown die-back. In severe cases, mortality occurs. For example, severe dewatering of channels of the braided Big Lost River in Idaho led to mortality of the narrowleaf cottonwood, P. angustifolia, and adjacent sandbar willows, Salix exigua Nutt., within 5 years, whereas riparian woodlands thrived along flowing channels nearby. The conservation and restoration of cottonwoods will rely on the provision of river flow regimes that satisfy these ecophysiological requirements for survival, growth and reproduction.
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