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Growth responses of five desert plants as influenced by biological soil crusts from a temperate desert, China

In almost all dryland systems, biological soil crusts (biocrusts) coexist alongside herbaceous and woody vegetation, creating landscape mosaics of vegetated and biocrusted patches. Results from past studies on the interaction between biocrusts and vascular plants have been contradictory. In the Gurbantunggut desert, a large temperate desert in northwestern China, well-developed lichen-dominated crusts dominate the areas at the base and between the sand dunes. We examined the influence of these lichen-dominated biocrusts on the germination, growth, biomass accumulation, and elemental content of five common plants in this desert: two shrubs ( Haloxylon persicum , Ephedra distachya ) and three herbaceous plants ( Ceratocarpus arenarius , Malcolmia africana and Lappula semiglabra ) under greenhouse conditions. The influence of biocrusts on seed germination was species-specific. Biocrusts did not affect percent germination in plants with smooth seeds, but inhibited germination of seeds with appendages that reduced or eliminated contact with the soil surface or prevented seeds from slipping into soil cracks. Once seeds had germinated, biocrusts had different influences on growth of shrub and herbaceous plants. The presence of biocrusts increased concentrations of nitrogen but did not affect phosphorus or potassium in tissue of all tested species, while the uptake of the other tested nutrients was species-specific. Our study showed that biocrusts can serve as a biological filter during seed germination and also can influence growth and elemental uptake. Therefore, they may be an important trigger for determining desert plant diversity and community composition in deserts.

Ecological Research

Impact of biological soil crusts and desert plants on soil microfaunal community composition

Carbon and nitrogen are supplied by a variety of sources in the desert food web; both vascular and non-vascular plants and cyanobacteria supply carbon, and cyanobacteria and plant-associated rhizosphere bacteria are sources of biological nitrogen fixation. The objective of this study was to compare the relative influence of vascular plants and biological soil crusts on desert soil nematode and protozoan abundance and community composition. In the first experiment, biological soil crusts were removed by physical trampling. Treatments with crust removed had fewer nematodes and a greater relative ratio of bacterivores to microphytophages than treatments with intact crust. However, protozoa composition was similar with or without the presence of crusts. In a second experiment, nematode community composition was characterized along a spatial gradient away from stems of grasses or shrubs. Although nematodes generally occurred in increasing abundance nearer to plant stems, some genera (such as the enrichment-type Panagrolaimus) increased disproportionately more than others (such as the stress-tolerant Acromoldavicus). We propose that the impact of biological soil crusts and desert plants on soil microfauna, as reflected in the community composition of microbivorous nematodes, is a combination of carbon input, microclimate amelioration, and altered soil hydrology. ?? Springer Science + Business Media B.V. 2009.

Plant and Soil

Dominant Sonoran Desert plant species have divergent phenological responses to climate change

The southwestern U.S. is a global hotspot of climate change. Models project that temperatures will continue to rise through the end of the 21st century, accompanied by significant changes to the hydrological cycle. Within the Sonoran Desert, a limited number of studies have documented climate change impacts on the phenology of native plant species. Much of this phenological work to understand climate change impacts to phenology builds on research conducted nearly three decades ago to define flowering triggers and developmental requirements for native keystone Sonoran Desert woody species. Here we expand on the drivers and explore recent phenological trends for six species using a unique 36-year observational data set. We use statistical models to determine which aspects of climate influence the probability of flowering, and how flowering time may respond to climate change. We move beyond traditional models of phenology by incorporating different metrics of moisture availability in addition to temperature, weather, and climate at several time scales, including daily, weekly, seasonal, and antecedent conditions. Our results provide evidence of a trend towards earlier flowering (on the order of 1–4 days per decade) for five of the six species analyzed, and no trend for one species. The species we evaluated had contrasting phenological responses to different aspects of climate, suggesting individualistic changes in phenology and the potential of divergent plant community flowering patterns under future climate change. Understanding recent changes in flowering phenology and their climatic triggers is important to anticipating whether plant species can attract pollinators, reproduce, and persist within the community under continued climate change.

Arizona

Succession of desert plants on debris flow terraces, Grand Canyon, Arizona, U.S.A.

Vegetation sampling on 11 debris flow terraces in Grand Canyon National Park, Arizona, U.S.A., showed that plant assemblages changed as age of surface increased. The terraces ranged in age from about 5 to about 3100 years. There were distinct differences among sites in the life history characteristics of the dominant plants. Young terraces (5-55 years) were dominated by short-lived plants that had high reproductive potential. Older surfaces were dominated by species with longer life-spans and lower reproductive potential. Density and cover of long-lived species increased with age of surface; for short-lived plants, density was inversely related to surface age. Species composition was also correlated with site age; however, location, exposure, and other factors ensured that no two debris flows supported identical mixtures of species. Succession on recent Grand Canyon debris flows is driven in part by life-history strategies, particularly life-span and seed dispersal traits, and also by climatic factors, especially those that control germination and establishment of the long-lived dominants.

Journal of Arid Environments

Effects of high fire frequency in creosote bush scrub vegetation of the Mojave Desert

Plant invasions can increase fire frequency in desert ecosystems where fires were historically infrequent. Although there are many resource management concerns associated with high frequency fire in deserts, fundamental effects on plant community characteristics remain largely unstudied. Here I describe the effects of fire frequency on creosote bush scrub vegetation in the Mojave Desert, USA. Biomass of the invasive annual grass Bromus rubens L. increased following fire, but did not increase further with additional fires. In contrast, density, cover and species richness of native perennial plants each decreased following fire and continued to decrease with subsequent fires, although not as dramatically as after the initial fire. Responses were similar 5 and 14 years post-fire, except that cover of Hymenoclea salsola Torr. & A. Gray and Achnatherum speciosa Trin. & Rupr. both increased in areas burnt once. These results suggest that control of B. rubens may be equally warranted after one, two or three fires, but revegetation of native perennial plants is most warranted following multiple fires. These results are valid within the scope of this study, which is defined as relatively short term vegetation responses (≤14 years) to short fire return intervals (6.3 and 7.3 years for the two and three fire frequency levels) within creosote bush scrub of the Mojave Desert.

Mojave Desert

Forecasting climate change impacts to plant community composition in the Sonoran Desert region

Hotter and drier conditions projected for the southwestern United States can have a large impact on the abundance and composition of long-lived desert plant species. We used long-term vegetation monitoring results from 39 large plots across four protected sites in the Sonoran Desert region to determine how plant species have responded to past climate variability. This cross-site analysis identified the plant species and functional types susceptible to climate change, the magnitude of their responses, and potential climate thresholds. In the relatively mesic mesquite savanna communities, perennial grasses declined with a decrease in annual precipitation, cacti increased, and there was a reversal of the Prosopis velutina expansion experienced in the 20th century in response to increasing mean annual temperature (MAT). In the more xeric Arizona Upland communities, the dominant leguminous tree, Cercidium microphyllum, declined on hillslopes, and the shrub Fouquieria splendens decreased, especially on south- and west-facing slopes in response to increasing MAT. In the most xeric shrublands, the codominant species Larrea tridentata and its hemiparasite Krameria grayi decreased with a decrease in cool season precipitation and increased aridity, respectively. This regional-scale assessment of plant species response to recent climate variability is critical for forecasting future shifts in plant community composition, structure, and productivity.

Sonoran Desert

Regional signatures of plant response to drought and elevated temperature across a desert ecosystem

The performance of many desert plant species in North America may decline with the warmer and drier conditions predicted by climate change models, thereby accelerating land degradation and reducing ecosystem productivity. We paired repeat measurements of plant canopy cover with climate at multiple sites across the Chihuahuan Desert over the last century to determine which plant species and functional types may be the most sensitive to climate change. We found that the dominant perennial grass, Bouteloua eriopoda, and species richness had nonlinear responses to summer precipitation, decreasing more in dry summers than increasing with wet summers. Dominant shrub species responded differently to the seasonality of precipitation and drought, but winter precipitation best explained changes in the cover of woody vegetation in upland grasslands and may contribute to woody-plant encroachment that is widespread throughout the southwestern United States and northern Mexico. Temperature explained additional variability of changes in cover of dominant and subdominant plant species. Using a novel empirically based approach we identified ‘‘climate pivot points’’ that were indicative of shifts from increasing to decreasing plant cover over a range of climatic conditions. Reductions in cover of annual and several perennial plant species, in addition to declines in species richness below the long-term summer precipitation mean across plant communities, indicate a decrease in the productivity for all but the most drought-tolerant perennial grasses and shrubs in the Chihuahuan Desert. Overall, our regional synthesis of long-term data provides a robust foundation for forecasting future shifts in the composition and structure of plant assemblages in the largest North American warm desert.

Ecology

Structural changes in vegetation coincident with annual grass invasion negatively impacts sprint velocity of small vertebrates

Sagebrush ecosystems in the intermountain west of the United States are being threatened by conversion to the non-native grass, cheatgrass ( Bromus tectorum ). The dramatic shift in the physical structure of vegetation coincident with cheatgrass invasion likely has negative impacts on animal communities, yet these structural impacts have not been well-studied. In a previous study, dense cheatgrass stems reduced sprint velocity for the flattened, wide-bodied desert horned lizard ( Phrynosoma platyrhinos ). Here, we asked if a decrease in sprint velocity due to cheatgrass impediment can be generalized to the suite of small vertebrates inhabiting the sagebrush ecosystems of western Utah. We evaluated sprint performance of the common rodent ( n = 3) and lizard ( n = 4) species on two raceway types, cheatgrass and no-cheatgrass, and hypothesized that body size, body shape, and form of movement are important factors influencing sprint velocity through dense cheatgrass stems. All species showed significant reductions in speed on cheatgrass versus no-cheatgrass raceways, with percent reduction greatest for larger, wider, or hopping organisms compared to smaller, more slender, or running organisms. Of concern, surveys for rodents and lizards at our study areas support a common pattern: lower abundances of small vertebrates, as well as a loss of rodent species richness, in areas infested with cheatgrass compared to intact, native sagebrush communities. By extension, we expect a negative impact on animal communities in other semi-arid regions experiencing dramatic shifts in vegetation structure upon invasion by non-native grasses that are capable of forming dense stands in the interspaces of native desert plants [e.g., Sonoran desert invaded by buffelgrass ( Pennisetum ciliare )].

Utah

Seed dispersal by specialist versus generalist foragers: The plant's perspective

I examined the seed dispersal ecology of the stem parasitic plant, desert mistletoe (Phoradendron californicum, Viscaceae), with the objectives of (1) determining the relative effectiveness of specialist and generalist foragers for seed dispersal, (2) determining the extent to which desert mistletoe fruiting characteristics correspond to those predicted for plants attracting specialist versus generalist foragers, and (3) examining the potential consequences of the observed dispersal strategy for mistletoe reproduction. Three species of birds, phainopepla, Gila woodpecker, and northern mockingbird, fed on desert mistletoe at my study site. The specialist, phainopepla, was the most abundant and the most likely to perch in host species, where defecated seeds had a greater probability of lodging in a site suitable for establishment. Gila woodpeckers, although abundant, spent little time in host plants, thus dooming most of the seeds they consumed. Mockingbirds may disperse a small number of seeds, but were abundant enough to consume only a small portion of the available fruits. As expected for plants attracting specialist frugivores, mistletoes produced fruits throughout the 6-month season in which phainopeplas reside in the Sonoran desert. Contrary to expectation, numbers of fruits produced far exceeded the amount that could be consumed by the frugivores at my study site. Fruit crop size was positively related to absolute fruit removal, but not to proportional removal at the scale of the entire study site. However, crop size was positively related to proportional removal within the neighborhood of mistletoes occupying an individual host tree. Frugivores were attracted to infected hosts, host attractiveness increased, although proportional removal of fruit declined, with number of female mistletoes. The observed dispersal ecology of desert mistletoe suggests the likelihood of increasingly clumped distributions of mistletoe plants, as more and more seeds are deposited on previously infected hosts, and increased density of mistletoes attract ever more visits by birds. I observed no decline in vigor, in terms of fruit production, within the levels of infestation at my study site. The seed dispersal strategy of desert mistletoe thus includes aspects of that expected both for plants dependent on specialists and those dependent on generalists. Fruits are available through the entire season to maintain the specialist. Production far exceeds that expected, but serves to attract the specialist within a neighborhood of vigorously fruiting conspecifics.

Oikos

Dust deposition effects on growth and physiology of the endangered Astragalus jaegerianus (Fabaceae)

Human expansion into the Mojave Desert is a significant threat to rare desert plants. While immediate habitat loss is often the greatest concern, rare plants situated near areas where soil surfaces experience frequent disturbance may be indirectly impacted when fine particulate dust accumulates on leaf surfaces. Remaining populations of the federally listed Astragalus jaegerianus (Lane Mountain milkvetch) occur on land open to expanding military activities and on adjacent public land with increasing recreational use. This study was initiated to determine whether dust accumulation could decrease the vigor and fitness of A. jaegerianus through reduced growth. Beginning in early May 2004, plants located on Bureau of Land Management (BLM) land were dusted bimonthly at canopy-level dust concentrations ranging from 0 to 32 g/m 2 , and physiology and growth were monitored until late June when plants senesced. The maximum experimental dust level simulates dust concentrations of Mojave Desert perennials neighboring military activities at a nearby army training center. Average shoot growth declined with increasing dust accumulation, but seasonal net photosynthesis increased. Further investigation of plants grown in a greenhouse supported similar trends. This pattern of greater net photosynthesis with increasing dust accumulation may be explained by higher leaf temperatures of dusted individuals. Ambient dust deposition measured in traps near field plants (May 2004–July 2004) ranged from 0.04–0.17 g/m 2 / d, which was well below the lowest level of dust on experimental plants (3.95 g/m 2 /d). With this low level of ambient deposition, we expect that A. jaegerianus plants in this population were not greatly affected by the dust they receive at the level of recreational use during the study.

Madroño

Effects of substrate disturbance on secondary plant succession; Mojave Desert, California.

(1) The effects of substrate disturbance on perennial plant succession in the Mojave Desert were assessed at three military camps abandoned for 40 years. (2) Soil compaction, removal of the top layer of soil, and alteration of drainage channel density caused significant changes in perennial plant cover, density, and relative species composition. (3) Long-lived species, predominantly Larrea tridentata , were dominant in all control areas but percentage cover and density were greatly reduced in areas where substrate alterations were significant. (4) Pioneer species such as Ambrosia dumosa and Hymenoclea salsola had percentage cover values similar to or greater than controls in most areas where substrate alterations were significant, and these species were dominant in the majority of disturbed areas. (5) Where substrate alterations were insignificant in disturbed areas at one camp, Larrea was the dominant species as in the control.

Journal of Applied Ecology

Interaction assessment: Rationale and a test using plants

A non-manipulative method for deriving empirical expressions of population growth parameters from simple field data is presented. The derived expressions can be used to assess the intensity and form of density dependence and interspecies interactions, and have potential for parameterizing more mechanistic models of population dynamics and for use in applied ecology, e.g. land management or environmental risk assessment. The method is based on an assertion of invariant expected fitness across occupied microhabitats. Hence, its success depends upon the degree to which that assertion holds. The assertion, as used here, is broadly applicable. Thus, the method can be expected to yield reliable results even in nonequilibrium communities. Here, we apply the method to data on six desert plant species. Expressions generated from data in one stand, in one year, successfully predict plant cover values in other stands and years. The predicted patterns of plant species interactions are discussed in the light of current knowledge and theorics of desert succession.

Evolutionary Ecology

Evidence against a Pleistocene desert refugium in the Lower Colorado River Basin

Aim The absence of Sonoran Desert plants in late Pleistocene-aged packrat middens has led to speculation that they survived glacial episodes either in refugia as intact associations (Clementsian community concept) or in dry microsites within chaparral or woodland according to individualistic species responses (Gleasonian community concept). To test these hypotheses, we developed a midden record from one likely refugium in north-eastern Baja California, Mexico. We also measured stomatal guard cell size in fossil leaves to further evaluate site-level individualistic responses of Larrea tridentata (creosote bush) ploidy races to climatic changes, including monsoonal history, over the late Quaternary. Location Sierra Ju&aacute;rez, Lower Colorado River Basin, north-eastern Baja California, Mexico. Methods Packrat ( Neotoma ) middens were collected from <300 m elevation on the eastern piedmont of the Sierra Ju&aacute;rez. Plant macrofossils and pollen were analysed from 50 dated middens, including determination of Larrea tridentata ploidy races. Results Pleistocene middens dating back to >55,000 cal. yr BP contained a mix of extralocal species characteristic of chaparral and pinyon&ndash;juniper&ndash;oak woodland, along with some modern desert elements. Many other desert taxa were absent during the Pleistocene, although most had arrived by the beginning of the Holocene 11,700 years ago. Main conclusions The assemblage of chaparral, woodland and select desert elements refutes the hypothesis that the Lower Colorado River Basin served as a late Pleistocene refugium for Sonoran Desert flora. The rapid arrival of most missing desert species by the early Holocene suggests they did not have far to migrate. They probably survived the last glacial period as smaller, disparate populations in dry microsites within chaparral and pinyon&ndash;juniper&ndash;oak woodlands. Diploid and tetraploid races of Larrea tridentata were present during the Pleistocene, but hexaploids did not appear until the mid-Holocene. This demonstrates that individualistic responses to climate involved genetic variants, in this case cytotypes, and not just species.

Baja California

Twentieth century demographic changes in cirio and cardón in Baja California, Mexico

Aim&ensp; Our purpose was to discern long-term large-scale patterns of survivorship and recruitment of two dominant, charismatic and useful desert plants, cirio [Fouquieriaceae; Fouquieria columnaris (Kell.) Kell. ex Curran] and card&oacute;n [Cactaceae; Pachycereus pringlei (S.Watson) Britton & Rose], and to evaluate the effects of physical and human factors. Location&ensp; The study included 77 sites distributed over c. 13,200 km 2 in the northern Vizca&iacute;no Region of the Sonoran Desert, state of Baja California, M&eacute;xico. Method&ensp; Censuses used identified individuals ( n = 3780 cirio and 2246 card&oacute;n) and were based on repeat photography. Time intervals between censuses ranged from 29 to 96 years, ending in 1996&ndash;2002. Earlier repetition (1963) of old photographs at 14 sites allowed analysis of temporal variation. The effect of elapsed time was modelled with Weibull functions for survivorship and linear functions for recruitment. To explain the residual variations, we tested categorical and continuous variables related to substrate, geomorphology, climate and geography. Human impacts were weighed with historical and quantitative analysis. Results&ensp; Elapsed time accounted for 45% of the variation of survivorship in cirio but only 35% in card&oacute;n. The fitted Weibull curves were used to estimate longevity: all-size cohorts of 1000 individuals would die out in 388 (cirio) and 723 years (card&oacute;n). Recruitment in cirio was linearly related to time ( r 2 = 0.71) and averaged < 0.003 year &minus;1 but was less than mortality at 90% of the sites. For card&oacute;n, recruitment averaged > 0.005 year &minus;1 , was linearly related to time ( r 2 = 0.66) and was less than mortality at only 33% of the sites. Vital rates were not strongly correlated between mid- and late-century but were mostly similar to the long-term regional rates. Removing the factor of elapsed time, the residual variations showed that survivorship was greater on sites protected from winds for cirio and was less on very coarse soils for card&oacute;n. Recruitment increased with latitude in cirio and was greater on southern exposures for both species. The residual variations were not clearly affected, at a regional scale, by other factors thought to be important, including elevation, distance to the Pacific Ocean, geology, slope gradient, soil stability, older vs. young alluvial soils, and soil Ca : Mg and Na : K ratios. Human impacts have been sporadic and heterogeneous but locally strong; our quantitative indices of accessibility did not show regionally significant effects. Blowdown by hurricanes is a sporadic cause of major mortality for cirio but not necessarily for card&oacute;n. Main conclusions&ensp; At our scales, effects of time outweigh those of substrate, and human impact was scant. Large patterns were pronounced: cirio experienced a prolonged, widespread decline in the 1900s, while card&oacute;n fluctuated in different directions and degree among local populations. Cirio was found to be inherently much shorter-lived than card&oacute;n. We also suggest that recruitment in cirio was low due to long periods with relatively dry winters that affected the entire region, while spatial heterogeneity of card&oacute;n recruitment seemed more related to the variation in summer rains.

Journal of Biogeography

Short seed longevity, variable germination conditions, and infrequent establishment events provide a narrow window for Yucca brevifolia (Agavaceae) recruitment

PREMISE OF THE STUDY: The future of long-lived stand-forming desert plants such as Yucca brevifolia (Joshua tree) has come into question in light of climate variation and landscape-scale disturbances such as wildfire. Understanding plant establishment dynamics is important for mitigating the impacts of disturbances and promoting revegetation. • METHODS: We placed Y. brevifolia seeds in shallow caches and manipulated granivore access, nurse shrub effects, and the season of cache placement to determine conditions for seed germination and seedling establishment. • KEY RESULTS: Greatest seedling emergence occurred during spring and summer, when increased soil moisture was accompanied by warm soil temperatures. Late winter-spring emergence for cached seeds was enhanced beneath shrub canopies, but seedling survival declined beneath shrubs as temperatures increased in spring. Germinability of seed remaining in the soil was reduced from 50-68% after 12 mo residence time in soil and declined to <3% after 40 mo. Following dispersal from parent plants, seeds are either removed by granivores or lose germinability, imposing substantial losses of potential germinants. • CONCLUSIONS: Specific germination and establishment requirements impose stringent limits on recruitment rates for Y. brevifolia. Coupled with infrequent seed availability, the return rates to prefire densities and demographic structure may require decades to centuries, especially in light of potential changes to regional desert climate in combination with the potential for fire recurrence. Demographic patterns are predicted to vary spatially in response to environmental variability that limits recruitment and may already be apparent among extant populations.

American Journal of Botany

Fire, climate and changing forests

A changing climate implies potential transformations in plant demography, communities, and disturbances such as wildfire and insect outbreaks. How do these dynamics play out in terrestrial ecosystems across scales of space and time? “Vegetation type conversion” (VTC) is a term used to describe abrupt and long-lasting changes in vegetation structure and composition due to various kinds of perturbations. For example, it has long been observed that fire-adapted ecosystems such as the California chaparral shrublands are readily replaced by non-native annual grasses when humans increase fire frequency. Similar effects are being observed in Sonoran Desert plant communities, where invasion by non-native buffelgrass (Cenchrus ciliare) exposes a fire-sensitive plant community to high-intensity fires, which are lethal to the long-lived iconic Sonoran flora such as Saguaro cactus (Carnegiea gigantea). While type conversions following severe disturbance have been observed for some time, the mechanisms that underlay such changes are only recently coming to light. Threshold behavior is of particular concern: as various climate indices pass critical values, some ecosystems may be reaching points of no return where recovery to the earlier state is no longer possible. A recent publication in the Proceedings of the National Academy of Sciences, U.S., has now revealed that increased fire severity impairs post-fire forest regeneration, setting the stage for potential large-scale type conversion of western conifer forests.

Nature Plants