To investigate the evolution of clinal variation in an invasive plant, we compared cold hardiness in the introduced saltcedar (Tamarix ramosissima, Tamarix chinensis, and hybrids) and the native plains cottonwood (Populus deltoides subsp. monilifera). In a shadehouse in Colorado (41°N), we grew plants collected along a latitudinal gradient in the central United States (29–48°N). On 17 occasions between September 2005 and June 2006, we determined killing temperatures using freeze-induced electrolyte leakage and direct observation. In midwinter, cottonwood survived cooling to −70°C, while saltcedar was killed at −33 to −47°C. Frost sensitivity, therefore, may limit northward expansion of saltcedar in North America. Both species demonstrated inherited latitudinal variation in cold hardiness. For example, from September through January killing temperatures for saltcedar from 29.18°N were 5–21°C higher than those for saltcedar from 47.60°N, and on September 26 and October 11, killing temperatures for cottonwood from 33.06°N were >43°C higher than those for cottonwood from 47.60°N. Analysis of nine microsatellite loci showed that southern saltcedars are more closely related to T. chinensis while northern plants are more closely related to T. ramosissima. Hybridization may have introduced the genetic variability necessary for rapid evolution of the cline in saltcedar cold hardiness.

To explore the roles of plasticity and genetic variation in the response to spatial and temporal climate variation, we established a common garden consisting of paired collections of native and introduced riparian trees sampled along a latitudinal gradient. The garden in Fort Collins, Colorado (latitude 40.6°N), included 681 native plains cottonwood (Populus deltoides subsp. monilifera) and introduced saltcedar (Tamarix ramosissima, T. chinensis and hybrids) collected from 15 sites at 29.2-47.6°N in the central United States. In the common garden both species showed latitudinal variation in fall, but not spring, leaf phenology, suggesting that the latitudinal gradient in fall phenology observed in the field results at least in part from inherited variation in the critical photoperiod, while the latitudinal gradient in spring phenology observed in the field is largely a plastic response to the temperature gradient. Populations from higher latitudes exhibited earlier bud set and leaf senescence. Cold hardiness varied latitudinally in both fall and spring for both species. For cottonwood, cold hardiness began earlier and ended later in northern than in southern populations. For saltcedar northern populations were hardier throughout the cold season than southern populations. Although cottonwood was hardier than saltcedar in midwinter, the reverse was true in late fall and early spring. The latitudinal variation in fall phenology and cold hardiness of saltcedar appears to have developed as a result of multiple introductions of genetically distinct populations, hybridization and natural selection in the 150 years since introduction.

Wild horses (Equus caballus) at Pryor Mountain were studied by direct observation from 1993 through 2007. All horses present were individually identifiable on the basis of coat coloration, head and leg markings, gender, and band associations. Of the 609 horses either present prior to foaling in 1993 or born since, ages were precisely known for density-dependent population regulation, as both population growth rate and survival rate were negatively correlated with population size from the previous year. These and other factors were not sufficient to stabilize the population during our period of study, however, as evidenced by the necessity for large removals in several years.

Abstract:We examined the response of first year saltcedar (Tamaria ramosissima) and plains cottonwood (Populus deltoides subsp, monilifera) seedlings to flooding in fall (25 days) and spring (28 days) using potgrown plants (12-18 individuals/26.5-1iter pot). Seedlings were initially counted in all pots prior to fall treatment. Survival was calculated as the proportion of seedlings in each pot still alive following spring treatment. Mean survival rates of seedlings flooded in fall (saitcedar = 0.8%, cottonwood = 20.8%, n = I4 pots) were lower compared to the spring flooding treatment (saltcedar -91.1%, cottonwood -92.2%, n = 13) and control (saltcedar = 93.9%, cottonwood = 98.7%, n = 14). We used multiple response permutation procedures to detect omnibus distributional differences in survival data (total tests = 9) because assumptions of normality and equal variance were not met. Survival distributions differed between saltcedar and cottonwood fall flooding groups (P < 0.0001) and between fall flooding and control groups for both species (P < O.O00l). No differences in survival distributions were detected between species or treatments for the control and spring treatment groups (P > 0.07). Smaller size and consequent lack of energy reserves may account for lower survival of saltcedar compared to cottonwood in the fall treatment and for lower survival of both species in the fall lreatment compared to the spring treatment. Fall flooding for controlling lirsl year saltcedar seedlings is suggested as a potentially useful technique in riparian habitat restoration and management in the southwestern United States.

The 4-year drawdown of Horsetooth Reservoir, Colorado, for dam maintenance, provides a case study analog of vegetation response on sediment that might be exposed from removal of a tall dam. Early vegetation recovery on the exposed reservoir bottom was a combination of (1) vegetation colonization on bare, moist substrates typical of riparian zones and reservoir sediment of shallow dams and (2) a shift in moisture status from mesic to the xeric conditions associated with the pre-impoundment upland position of most of the drawdown zone. Plant communities changed rapidly during the first four years of exposure, but were still substantially different from the background upland plant community. Predictions from the recruitment box model about the locations of Populus deltoides subsp. monilifera (plains cottonwood) seedlings relative to the water surface were qualitatively confirmed with respect to optimum locations. However, the extreme vertical range of water surface elevations produced cottonwood seed regeneration well outside the predicted limits of drawdown rate and height above late summer stage. The establishment and survival of cottonwood at high elevations and the differences between the upland plant community and the community that had developed after four years of exposure suggest that vegetation recovery following tall dam removal will follow a trajectory very different from a simple reversal of the response to dam construction, involving not only long time scales of establishment and growth of upland vegetation, but also possibly decades of persistence of legacy vegetation established during the reservoir to upland transition.