Experimental manipulations of precipitation seasonality: effects on oak ( Quercus ) seedling demography and physiology

Predicted changes in regional precipitation patterns and soil moisture caused by anthropogenic trace gas emissions may affect the distribution and abundance of woody plants in arid and semiarid regions. To test the response of woody plants to potential changes in precipitation regimes, we manipulated summer and winter precipitation on plots that contained seedlings of Quercus emoryi Torr. (Emory oak), the dominant tree in oak savannas of the southwestern United States. Throughout the growing season, we monitored seedling survival and physiology (predawn leaf water potential, midday instantaneous gas exchange, and leaf carbon isotope discrimination). Seedling survival and physiological performance differed little between treatments, which embodied 50% changes to quantities of summer and winter precipitation, and encompassed a continuum of precipitation from 359 mm · year -1 to 846 mm · year -1 . However, survival and physiological performance of seedlings were negatively impacted by seasonal environmental conditions common to all treatments, especially during the annual pre-'monsoon' drought. Seedling predawn leaf water potentials, net CO 2 assimilation, and stomatal conductance indicate that growing conditions for Q. emoryi seedlings at this site are generally restricted to periods with adequate soil moisture (i.e., April and August). Results contrast with an assumption implicit to the ""two-layer"" soil water resource partitioning hypothesis that woody plants in all life history stages are more dependent upon winter than summer precipitation. In fact, summer precipitation appears more important than winter precipitation for Q. emoryi seedling recruitment and growth.     

Tree growth and regeneration response to climate and stream fow in a species-rich southwestern riparian forest

We studied the influence of climate variables, stream flow, and topography on regeneration and growth of several riparian tree species ( Acer negundo, Alnus oblongifolia, Fraxinus velutina, Juglans major, Platanus wrightii, Populus fremontii, Salix spp.) at an unregulated perennial stream, West Clear Creek, in central Arizona. A pulse of seedling regeneration occurred for Alnus, Fraxinus, Platanus, Populus , and Salix in 1995 and 1996 following high winter and spring surface flows in 1993 and high spring surface flow in 1995. In contrast, little regeneration occurred for Acer and Juglans under these conditions. Most seedlings occurred at the active channel topographic location, and few seedlings occurred at abandoned channel, gravel-boulder bar, and bench locations. Relationships between environmental variables and annual radial stem growth varied among species and between constrained and unconstrained reaches. High spring or winter surface flows were negatively related to growth of Acer, Alnus, and Platanus , whereas high spring surface flow was positively related to growth of Fraxinus . Positive relationships between precipitation and growth occurred only for Fraxinus and Juglans , suggesting greater use of surface soil water by these species. Annual radial growth was high for Platanus and Alnus, medium for Acer and Fraxinus, and low for Juglans . Overall, the tree species in our study responded individually, rather than collectively, in regeneration and growth to changes in stream flow and climatic variables.     

Seed production and seedling establishment of a Southwest riparian tree, Arizona walnut ( Juglans major )

A four-year study of five populations has revealed influences on seed production and seedling establishment of the Southwest riparian tree Juglans major . Germination is abundant after production of large seed crops (masts), but masts are produced infrequently. Within years, germination is stimulated by summer rains, enabling seedlings to establish on riparian terraces as well as streambanks. Traits such as capacity for dormancy during summer drought allow some seedlings to survive on terraces, but abundant rainfall is essential for high rates of seedling success. Ranges of moisture tolerance vary among seeds collected from different populations, suggesting that ecotypes may exist between riparian sites with dissimilar moisture regimes. Population-based differences are associated, in part, with differences in seed size.     

Flood flows and population dynamics of Arizona sycamore ( Platanus wrightii )

Platanus wrightii is a pioneer tree species of warm-temperate riparian deciduous forests in southwestern United States and northern Mexico. Dendrochronological analysis of populations in central and southern Arizona indicated that P. wrightii seedlings establish episodically. Long intervals (10-40 years) elapsed with no apparent establishment. Seedling establishment years were positively associated with winter flood size and annual flow rate, and weakly negatively associated with summer flood size. Large floods sometimes preceded multi-year establishment waves. During the past 2 decades an abundance of winter floods and very wet springs has allowed frequent establishment of P. wrightii . For example, plants established at many sites during the winter flood years of 1993 and 1995, particularly along channels scoured and widened by flood waters. Platanus wrightii also reproduces asexually. Ramets were more abundant than genets in all populations, and ramets established more frequently than seedlings. Ramet density (mean number per genet) varied widely between populations, from 2 (Huachuca Canyon) to 9 (Haunted Canyon), and increased with tree size within 4 of 9 populations. Population size structure varied across a gradient of watershed area. Populations along streams draining the largest watersheds had an abundance of small trees (mean trunk diameter of P. wrightii is influenced by temporal as well as spatial differences in stream flow regimes.     

Dynamics of a dwarf bear-poppy ( Arctomecon humilis ) population over a sixteen-year period

A population of the dwarf bear-poppy ( Arctomecon humilis Coville, Papaveraceae) at Red Bluff, Washington County, Utah, was monitored twice annually between 1987 and 2002. This is a narrowly endemic, gypsophilous species that has been formally listed as endangered since 1979. During the 16 years of observation, density of this species has fluctuated between 3 and 1336 individuals on the 0.07-ha monitoring plot. Moderate to large recruitments of seedlings occurred in 1992, 1995, and 2001. Seedling recruitments from a large, long-lived seed bank are triggered by abundant precipitation during the February-April period. At least 5.0 cm of rainfall is required during that interval to produce any seedlings. Seedlings experienced considerable mortality in the 1st few months of life in all observed cases. The average seedling initiated in the very large recruitment event of 1992 survived for only 2.6 years. Seedlings in that cohort that were alive 1 year after germination had an average longevity of 4.6 years. None of the seedlings that emerged in 1992 were still alive in October 2002. Mortality in this species was poorly correlated with fluctuations in precipitation or temperature. No epidemics of parasites or herbivores were observed. Mortality in the species appears to be caused by a variety of factors acting over a cohort's lifetime.     

Plant age/size distributions in black sagebrush ( Artemisia nova ): effects on community structure

The demography of black sagebrush ( Artemisia nova Nelson) was investigated in the Buckskin Mountains of western Nevada to determine patterns of stand renewal in sagebrush communities currently free from wildfires. Biomass sampling was conducted to develop growth classes that reflected apparent age of the shrubs. The density of black sagebrush plants was twice that of basin big sagebrush ( A. tridentata ssp. tridentata Nutt.) in adjacent communities on contrasting soils (2.2 versus 1.1 plants per m 2 ). Black sagebrush accumulated only 75% as much woody biomass as big sagebrush. Regression equations were developed and tested for predicting total woody biomass, current annual growth (CAG), and leaf weight of black sagebrush plants. Apparent age classes were developed both for the black sagebrush plants and the sub-canopy mounds on which they grew. Discriminant analysis was used to test this classification system. Plant succession, apparently controlled by nitrate content of the surface soil, appeared to eliminate the successful establishment of black sagebrush seedlings on the mounds. After the shrubs die, the mounds eventually deflate. We propose that mounds reform around shrub seedlings, but because seedling establishment is so rare in these communities, this could not be verified.     

Food of larval tui chubs, Gila bicolor, in Pyramid Lake, Nevada

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Vegetation, soils, and hydrogeomorphology of riparian patch types of a dryland river

From a landscape perspective, riparian corridors can be viewed as mosaics of vegetation patches. We delineated 10 patch types within the floodplain of the San Pedro River (Arizona) on the basis of physiognomy, dominant overstory species, and tree size class; and we assessed differences in hydrogeomorphology, vegetation structure, plant species richness, and soil chemistry and texture. Patches of tamarisk ( Tamarix ), an introduced species, fell within the continuum of variation shown by other patch types in the landscape mosaic. Among the tree-dominated types, cottonwood-willow ( Populus-Salix ) and tamarisk patches were inundated more frequently than mesquite ( Prosopis ) patches, while cottonwood-willow patches had shallower groundwater than tamarisk or mesquite patches. Due to the wetter conditions, cottonwood-willow patches had a high relative abundance of wetland and exotic species in the understory. Tamarisk patches and wet shrublands ( Baccharis salicifolia—Salix exigua ) had high woody stem densities while cottonwood-willow patches had dense canopy cover. In association with differences in canopy cover, cottonwood-willow patches had low herbaceous species richness but high woody species richness, while tamarisk patches had high herbaceous and low woody species richness. Soil electrical conductivity, silt content, organic matter content, and available phosphorus increased from young to old stands of both tamarisk and cottonwood-willow, often resulting in greater differences between patches of different size/age class than between patches with different dominant species. Surface soil salinity (electrical conductivity) was low in all patches, including those dominated by tamarisk. Nitrate was abundant in soils of tamarisk patches (perhaps reflecting their high clay content) and wet shrubland patches. Dry shrublands ( Hymenoclea-Ericameria ) and wet shrublands were similar to young forest patches in having coarse soils with little organic matter.     

Large-scale effects on bird assemblages in desert grasslands

During winter 1996–1997 and summer 1997, we surveyed birds at 26 study sites in desert grasslands of Arizona and New Mexico to determine tolerance of birds to variability in plant composition and structure. The relationship between bird abundance and vegetative characteristics might be an important consideration in the development of management and restoration plans. Of the 49 bird species we observed, we examined 13 and 16 species in detail for winter and summer seasons, respectively. A noticeable shift in species composition occurred between 3% and 10% woody plant cover. During winter, Chestnut-collared Longspurs ( Calcarius ornatus ) and Horned Larks ( Eremophila alpestris ) showed significant positive relationships with grass cover and negative relationships with woody plant cover, occurring in greatest numbers where woody cover was <1%. The richness of sparrow species during winter was highest among sites with 6% to 15% woody plant cover. In addition to showing significant positive relationships with woody cover, Vesper Sparrows ( Pooecetes gramineus ), Black-throated Sparrows ( Amphispiza bilineata ), Brewer’s Sparrows ( Spizella breweri ), and Chipping Sparrows ( S. passerina ) showed significant positive relationships with shrub species richness. Our results showed that reductions of woody plant cover below 3% in combination with the presence of native grasses could substantially enhance the presence of several bird species. Maintenance of woody cover between 6% and 15% could increase use by a suite of other species, especially wintering sparrows.     

Seed dispersal, seed entrapment, and seedling recruitment in a temporally variable desert playa

Natural establishment of seedlings in desert playas with temporally variable precipitation hinges on many factors, including seed production, seed dispersal, seed entrapment, seed germination, and seedling survival. We investigated natural seed dispersal patterns and the effects of surface texture, wind barriers, resource availability (water and nutrients), and protection from herbivory on seedling establishment in a desert playa. We hypothesized that the seed rain would be consistent throughout the year and that seedling establishment would improve in resource-amended plots with barriers to wind dispersal. Contrary to our hypotheses, seed flux peaked seasonally during the winter, and fertilization had no consistent effect on seedling establishment. Seed availability for dispersal correlated with precipitation in the previous water year, whereas seedling recruitment was greatest when current-year precipitation during the spring germination and summer seedling growth periods was high. Gravel and barrier surface treatments contained more surviving seedlings than other surface treatments. Comparison with plots located outside of herbivore exclosures, however, showed that greater seedling presence in gravel plots may be due somewhat to protection from herbivory provided by the gravel, rather than simply to the greater seed-trapping quality of the gravel. With abundant seed availability, application of surface treatments, like coarse gravel, combined with increased seasonal water availability could lead to improved shrub establishment from seed in desert playas.     

Seedling size and survival for Chrysothamnus nauseosus

Seedling size and survival in relation to summer drought were examined for Chrysothamnus nauseosus growing under field and greenhouse conditions. In the field, summer survival rates were less than 2% annually for the three years monitored. The effect of initial seedling height on subsequent survival was examined in both the field and greenhouse by grouping seedlings into live and dead categories on each census date and comparing initial heights for seedlings in these categories. For a majority of the census dates, the initial height of surviving seedlings was greater than the initial height of those that subsequently died (significant differences ranged from 1 to 8 mm), indicating that seedlings that were taller at the initiation of the drought period had a higher probability of survival. In the greenhouse, taller seedlings had greater shoot and root biomass and rooting depth. Seedlings that are larger (i.e., taller and have greater aboveground biomass) in late spring appear to have a higher probability of surviving summer drought due to greater rooting depth and hence increased access to moisture in deeper soil layers. Seed availability and safe sites for germination were probably not limiting since large numbers of seedlings successfully germinated in a patchy pattern during the study period. Seedling size and probability of survival were not related to either seedling density or the distance to nearest seedling neighbor. Survival through summer drought appears to be the main limitation to seedling recruitment in this population.     

Mapping aquatic faunal exchanges across the Sunda shelf,South-East Asia,using distributional and genetic data sets from the cyprinid fish Barbodes gonionotus (Bleeker, 1850)

The islands of Sumatra, Java and Borneo and mainland South-East Asia are encompassed by a shallow marine shelf, the Sunda shelf. The striking faunal similarities between countries bordering the Sunda shelf have been attributed to faunal exchanges across the shelf during Pleistocene glacial maxima when sea levels fell by up to 120 m and the shelf was exposed and drained by large river systems which connected many of today's rivers. The widespread distribution of fish species reflecting extensive faunal exchange across the area is predicted to be observed within species through the broad geographic distribution of genetic lineages. Here I report a discontinuous species distribution across the region for the cyprinid fish, Barbodes gonionotus. Mitochondrial DNA control region sequence and nuclear locus data from populations sampled across the species range, suggest that the present species distribution largely reflects faunal exchanges early in the Pleistocene and that these two regional population groupings have subsequently diverged in isolation. However, the mtDNA data present some evidence for limited genetic dispersal during more recent Pleistocene low sea levels. These results broadly confirm previous work from a catfish species and suggest that freshwater faunal exchanges may have been limited across the region during the Pleistocene despite extended periods during which the Sunda shelf was exposed.     

Notes on significant collections and additions to the flora of Glen Canyon National Recreation Area, Utah and Arizona, between 1992 and 2004

Symphyotrichum expansum (Puepp ex Spreng.) Nesom is reported new to Utah from the Escalante River drainage. A major range extension is reported for Aralia racemosa L. in the Escalante drainage, and additional populations are reported of the rare species Imperata brevifolia Vasey in Utah, including the 1st record for the Grand Staircase--Escalante National Monument. Heterotheca grandiflora Nutt. is reported new to north central Arizona. New locations and notes on an additional 22 rare species in Glen Canyon National Recreation Area are listed.     

Checklist of the mosses of Grand Teton National Park and Teton County, Wyoming

Symphyotrichum expansum (Puepp ex Spreng.) Nesom is reported new to Utah from the Escalante River drainage. A major range extension is reported for Aralia racemosa L. in the Escalante drainage, and additional populations are reported of the rare species Imperata brevifolia Vasey in Utah, including the 1st record for the Grand Staircase--Escalante National Monument. Heterotheca grandiflora Nutt. is reported new to north central Arizona. New locations and notes on an additional 22 rare species in Glen Canyon National Recreation Area are listed.     

Historic expansion of Juniperus occidentalis (western juniper) in southeastern Oregon

The chronology of Juniperus occidentalis (western juniper) expansion in eastern Oregon, the effect of plant canopy and interspace on J. occidentalis seedling establishment and growth rates, and the age of J. occidentalis maximum reproductive potential were determined. Measurements were recorded in twenty-two 0.4-ha plots established in sagebrush-grassland communities and six 0.1-ha plots in Populus tremuloides (quaking aspen) communities. J. occidentalis began increasing during the 1880s in stands containing > 130 yr old. Relatively steady establishment ensued into the 1950s and then began to progress at a geometric rate in the 19602. J. occidentalis encroachment into aspen stands began between 1910 and 1920. The largest proportion of juvenile trees established beneath Artemisia species in sagebrush-grassland communities. J. occidentalis trees appeared to reach full reproductive potential at > 50 yr of age. The ratio of male:female trees increased from 1.7 in scattered J. occidentalis stands to 3.8 in closed stands. The initiation of J. occidentalis encroachment during the late 1800s coincides with optimal climatic conditions for Juniperus berry production and establishment, reduced fire-return intervals, and heavy livestock grazing. The accelerated increase in J. occidentalis expansion since 1960 may be due to the continued absence of fire, abundant woody plant cover, and the large increase in J. occidentalis seed production.     

Distributional information on birds from egg sets collected by Henry Rogers Durkee in 1870 in southwestern Wyoming

Henry Rogers Durkee collected 74 egg sets of 27 avian species in 1870 at Gilmer, Uinta County, in southwestern Wyoming. Despite the paucity of documented breeding evidence from this region during the late 19th and early 20th centuries, his material at the Smithsonian Institution was generally overlooked and has never been critically examined. Durkee's egg sets included 5 species (Sandhill Crane, Grus candensis ; Northern Rough-winged Swallow, Stelgidoteryx serripennis ; Grasshopper Sparrow, Ammodramus savannarum ; Fox Sparrow, Passerella iliaca ; Cassin's Finch, Carpodacus cassinii ) whose breeding distribution was then poorly known, 25 to more than 70 years before nests and eggs were otherwise documented in Wyoming. Durkee also collected complete egg sets of Lark Bunting ( Calamospiza melanocorys ) over 70 years before breeding was confirmed at other peripheral locations in Wyoming. In addition, Durkee's incomplete egg sets of Grasshopper Sparrow were the 1st western subspecies ( A. s. perpallidus ); his complete egg set and nest of the Fox Sparrow collected at Gilmer constituted the 2nd locality for the species or species group ( P. i. schistacea ). Although the number of egg sets Durkee collected is modest, he made a meaningful contribution to the early history of avifaunal exploration in Wyoming.     

Competition Between Adult and Seedling Shrubs of Ambrosia dumosa in the Mojave Desert, Nevada

Seeds of the perennial shrub Ambrosia dumosa germinated in abundance following 11 days of rain during August 1983 at a study site in the northern Mojave Desert. Seedling establishment, growth, and reproduction were observed in natural vegetation and in an area that had been previously cleared of vegetation. For 5,527 A. dumosa seedlings, percent survival in April 1986 averaged 3% in the undisturbed vegetation and 58% in the denuded area. Seedlings occupying the cleared area had grown to sizes up to 0. 1 m 3 m by October 1984; some produced flowers and fruit in the spring of 1985. Surviving seedlings in the undisturbed vegetation were all smaller than 0.001 m 3 and did not reproduce. These pronounced differences in growth, survival, and reproduction associated with the presence or absence of adult shrubs demonstrated an intense competition that is incompatible with indications of mild competition from nearest-neighbor analyses. I therefore hypothesize that competition for water occurred, not by competition for water in two dimensions but by rapid use of a common resource, as if several people were drinking with straws from a common cup. This temporal mechanism would strongly favor adults over seedlings.     

The successional status of Cupressus arizonica

Several investigations (size-class analysis, age-determination inquiries, and germination tests) suggest that Cupressus arizonica of southeastern Arizona is a pioneer species. The tree requires disturbance to remove or reduce soil litter, which otherwise inhibits the reproduction of the species. Reduction of light intensity caused by canopy closure appears to be less important than litter accumulation in restricting C. arizonica reproduction. Following disturbance, successful establishment of seedlings may occur over an extended period (50 to 100 years), as litter gradually accumulates. The absence of C. arizonica seedlings in present populations suggest that fire suppression policies on federal lands where C. arizonica occurs have altered fire frequency, and consequently have fostered a short-term reduction in C. arizonica establishment. Only in floodplain environments, where flooding disturbs the soil surface, has much reproduction occurred in recent years. The long-term population pattern of C. arizonica appears stable, due to the great longevity of the species.     

Cryptogamic soil crusts: seasonal variation in algal populations in the Tintic Mountains, Juab County, Utah

The soil algae in the Tintic Mountains, Juab County, Utah, was studied over a one-year period in 1982 and 1983. Fluorescence microscopy was used to measure algal density in samples directly from the field. A total of 30 algal taxa was observed, blue-green algae being most abundant both in terms of density and number of species. Algal density showed peaks in late fall and late spring. Minima were present in September 1982 and July 1983. Several weak correlations between algal density and climatic data existed. In general algae correlated positively with precipitation and negatively with temperature. A combination of low precipitation and hot temperatures was likely responsible for the low density observed in July. The Chrysophyta followed slightly different trends than the other algal groups, having minima in early October 1982 and late August 1983. Field observations indicated that the degree of algal crusting varied noticeably over a period of one year, with highest abundance of hummocking in spring. Heavy summer thunderstorms destroyed algal crusting during July and August of 1983, though absolute density of algae increased during this time in response to the extra moisture.