Above- and belowground responses to nitrogen addition in a Chihuahuan Desert grassland

Increased available soil nitrogen can increase biomass, lower species richness, alter soil chemistry and modify community structure in herbaceous ecosystems worldwide. Although increased nitrogen availability typically increases aboveground production and decreases species richness in mesic systems, the impacts of nitrogen additions on semiarid ecosystems remain unclear. To determine how a semiarid grassland responds to increased nitrogen availability, we examined plant community structure and above- and belowground net primary production in response to long-term nitrogen addition in a desert grassland in central New Mexico, USA. Plots were fertilized annually (10 g N m⁻²) since 1995 and NPP measured from 2004 to 2009. Differences in aboveground NPP between fertilized and control treatments occurred in 2004 following a prescribed fire and in 2006 when precipitation was double the long-term average during the summer monsoon. Presumably, nitrogen only became limiting once drought stress was alleviated. Belowground NPP was also related to precipitation, and greatest root growth occurred the year following the wettest summer, decreasing gradually thereafter. Belowground production was unrelated to aboveground production within years and unrelated to nitrogen enrichment. Species richness changed between years in response to seasonal precipitation variability, but was not altered by nitrogen addition. Community structure did respond to nitrogen fertilization primarily through increased abundance of two dominant perennial grasses. These results were contrary to most nitrogen addition studies that find increased biomass and decreased species richness with nitrogen fertilization. Therefore, factors other than nitrogen deposition, such as fire or drought, may play a stronger role in shaping semiarid grassland communities than soil fertility.     

Precipitation magnitude and timing differentially affect species richness and plant density in the sotol grassland of the Chihuahuan Desert

Arid and semi-arid environments are dynamic ecosystems with highly variable precipitation, resulting in diverse plant communities. Changes in the timing and magnitude of precipitation due to global climate change may further alter plant community composition in desert regions. In this study, we assessed changes in species richness and plant density at the community, functional group, and species level in response to variation in the magnitude of natural seasonal precipitation and 25% increases in seasonal precipitation [e.g., supplemental watering in summer, winter, or summer and winter (SW)] over a 5-year period in a sotol grassland in the Chihuahuan Desert. Community species richness was higher with increasing winter precipitation while community plant density increased with greater amounts of winter and summer precipitation, suggesting winter precipitation was important for species recruitment and summer precipitation promoted growth of existing species. Herb and grass density increased with increasing winter and summer precipitation, but only grass density showed a significant response to supplemental watering treatments (SW treatment plots had higher grass density). Shrubs and succulents did not exhibit changes in richness or density in response to natural or supplemental precipitation. In this 5-year study, changes in community species richness and density were driven by responses of herb and grass species that favored more frequent small precipitation events, shorter inter-pulse duration, and higher soil moisture. However, due to the long life spans of the shrub and succulent species within this community, 5 years may be insufficient to accurately evaluate their response to variable timing and magnitude of precipitation in this mid-elevation grassland.     

Effects of kangaroo rat exclusion on vegetation structure and plant species diversity in the Chihuahuan Desert

Long-term (1977–90) experimental exclusion of three species of kangaroo rats from study plots in the Chihuahuan Desert resulted in significant increases in abundance of a tall annual grass (Aristida adscensionis) and a perennial bunch grass (Eragrostis lehmanniana). This change in the vegetative cover affected use of these plots by several other rodent species and by foraging birds. The mechanism producing this change probably involves a combination of decreased soil disturbance and reduced predation on large-sized seeds when kangaroo rats are absent. Species diversity of summer annual dicots was greater on plots where kangaroo rats were present, as predicted by keystone predator models. However, it is not clear whether this was caused directly by activities of the kangaroo rats or indirectly as a consequence of the increase in grass cover. No experimental effect on species diversity of winter annual dicots was detected. Our study site was located in a natural transition between desert scrub and grassland, where abiotic conditions and the effects of organisms may be particularly influential in determining the structure and composition of vegetation. Under these conditions kangaroo rats have a dramatic effect on plant cover and species composition.     

Effects of long-term rainfall variability on evapotranspiration and soil water distribution in the Chihuahuan Desert: A modeling analysis

We used the patch arid land simulator (PALS-FT) – a simple, mechanistic ecosystem model – to explore long-term variation in evapotranspiration (ET) as a function of variability in rainfall and plant functional type (FT) at a warm desert site in southern New Mexico. PALS-FT predicts soil evaporation and plant transpiration of a canopy composed of five principal plant FTs: annuals, perennial forbs, C₄ grasses, sub-shrubs, and evergreen shrubs. For each FT, the fractional contribution to transpiration depends upon phenological activity and cover as well as daily leaf stomatal conductance, which is a function of plant water potential, calculated from root-weighted soil water potential in six soil layers. Simulations of water loss from two plant community types (grass- vs. shrub-dominated) were carried out for the Jornada Basin, New Mexico, using 100 years of daily precipitation data (1891–1990). In order to emphasize variability associated with rainfall and fundamental differences in FT composition between communities, the seasonal patterns cover of perennials were held constant from year to year. Because the relative amount of year to year cover of winter and summer annual species is highly variable in this ecosystem, we examined their influence on model predictions of ET by allowing their cover to be variable, fixed, or absent.Over the entire 100-yr period, total annual ET is highly correlated with total annual rainfall in both community types, although T and E alone are less strongly correlated with rainfall, and variation in transpiration is nearly 3 times greater than evaporation and 2 times greater than variation in rainfall (CV of rainfall = 35%). Water use shows a relatively high similarity between the grass- and shrub-dominated communities, with a 100-yr average T/ET of 34% for both communities. However, based on a year-by-year comparison between communities, T/ET was significantly greater in the grass-dominated community, reflecting the fact that over the long term more than half of the rain occurs in the summer and is used slightly more efficiently (T¿E) by the C₄-grass community than the shrub community, although we found some rainfall patterns that resulted in much greater T/ET in the shrub community in a given year. Percent of water lost as transpiration (T/ET) suggests that while there is a general trend toward increased T/ET with rainfall in both community types, T/ET is extremely variable over the 100-yr simulation, especially for normal and below normal amounts of rainfall (T/ET values range from 1 to 58% for the grass-dominated site and 6 to 60% for the shrub-dominated site).These predictions suggest that because of the relatively shallow distribution of soil water, there is little opportunity for vertical partitioning of the soil water resource by differential rooting depths of the plant FTs, in contrast to the two-layer hypothesis of Walter (1971). However, functional types may avoid competition by keying on particular `windows' of moisture availability via differences in phenologies. We found very little differences in average, long-term model predictions of T, E, and ET when annual plant cover was variable, fixed, or absent. The results of our simulations help reconcile some of the disparate conclusions drawn from experimental studies about the relative contribution of transpiration vs. evaporation to total evapotranspiration, primarily by revealing the great year-to-year variability that is possible.     

Modifying rainfall patterns in a Mediterranean shrubland: system design,plant responses,and experimental burning

Global warming is projected to increase the frequency and intensity of droughts in the Mediterranean region, as well as the occurrence of large fires. Understanding the interactions between drought, fire and plant responses is therefore important. In this study, we present an experiment in which rainfall patterns were modified to simulate various levels of drought in a Mediterranean shrubland of central Spain dominated by Cistus ladanifer, Erica arborea and Phillyrea angustifolia. A system composed of automatic rainout shelters with an irrigation facility was used. It was designed to be applied in vegetation 2 m tall, treat relatively large areas (36 m²), and be quickly dismantled to perform experimental burning and reassembled back again. Twenty plots were subjected to four rainfall treatments from early spring: natural rainfall, long-term average rainfall (2 months drought), moderate drought (25% reduction from long-term rainfall, 5 months drought) and severe drought (45% reduction, 7 months drought). The plots were burned in late summer, without interfering with rainfall manipulations. Results indicated that rainfall manipulations caused differences in soil moisture among treatments, leading to reduced water availability and growth of C. ladanifer and E. arborea in the drought treatments. However, P. angustifolia was not affected by the manipulations. Rainout shelters had a negligible impact on plot microenvironment. Experimental burns were of high fire intensity, without differences among treatments. Our system provides a tool to study the combined effects of drought and fire on vegetation, which is important to assess the threats posed by climate change in Mediterranean environments.     

Effects of nitrogen amendment on annual plants in the Chihuahuan Desert

Effects of nitrogen amendments on spring annual plant distribution, primary production, and species diversity in a semi-arid environment were studied. The ecological responses of spring annual plant species to nitrogen differed between species, and between sites. The distribution of each species in a control transect was wider than in a nitrogen-treated transect. Annual plant species diversity at each station in the control transect was higher than that of the nitrogen-treated transect. The lower site of the Lower Basin Slope areas had the highest species diversity (0.94 for the control and 0.84 for the nitrogen-treated), and the Upper Basin Slope area, shrub vegetation zone, had the lowest species diversity (0.27 for the control and 0.05 for the nitrogen-treated) in both transects. Inorganic nitrogen in the nitrogen-treated transect soils was consistently higher than that in the control transect soils; however, the former showed more fluctuation from station to station than the latter. Above-ground biomass of spring annual plants in the nitrogen-treated transect was consistently higher than that in the control at each station. The maximum above-ground biomass in the control and nitrogen-treated transect was 24.4±4.4 gm⁻² and 61.2±10.6 gm⁻², respectively. Variations in above-ground biomass along the transect did not parallel with the variation in inorganic nitrogen in soils and species diversity.     

Elevated UV-B radiation effects on experimental grassland communities

Experimental grassland communities (turves) were exposed to supplemental levels of UV-B radiation (280–315 nm) at an outdoor facility, under treatment arrays of cellulose diacetate-filtered fluorescent lamps which also produce UV-A radiation (315–400 nm). Control treatments consisted of arrays of polyester-filtered lamps, which allowed for exposure to UV-A radiation alone, and arrays of unenergized lamps allowing for exposure to ambient levels of solar radiation.     

Microhabitat differentiation in Chiuhuahuan Desert plant communities

The effects of microhabitat differentiation on small-scale plant community structure in the Chiuhuahuan Desert were studied using multivariate analysis. The results showed that microhabitats (i.e., kangaroo rat mounds, ant mounds, shrubs, half-shrubs, and open areas) played a critical role in structuring small-scale plant community structure and maintaining species diversity. Annual plants were much more sensitvive to the presence of differentiated microhabitats than perennials and winter annuals exhibited stronger microhabitat perferences than summer annuals. Species diversity was highest on ant mounds while open areas supported the lowest diversity during both winter and summer. Biomass was highest in the shrub habitats followed by kangaroo rat mounds, ant mounds, half-shrubs, and open areas. Much of the diversity of these plants could be explained by the individualistic responses of species to the biotic effect of other plants or to disturbance by animals, or individualistic responses of species to differences in microenvironments.     

Sensitivity of three grassland communities to simulated extreme temperature and rainfall events

Three grassland communities in New Zealand with differing climates and proportions of C3 and C4 species were subjected to one‐off extreme heating (eight hours at 52.5°C) and rainfall (the equivalent of 100 mm) events. A novel experimental technique using portable computer‐controlled chambers simulated the extreme heating events. The productive, moist C3/C4 community was the most sensitive to the extreme events in terms of short‐term community composition compared with a dry C3/C4 community or an exclusively C3 community. An extreme heating event caused the greatest change to plant community species abundance by favouring the expansion of C4 species relative to C3 species, shifting C4 species abundance from 43% up to 84% at the productive, moist site. This was observed both in the presence and absence of added water. In the absence of C4 species, heating reduced community productivity by over 60%. The short‐term shifts in the abundance of C3 and C4 species in response to the single extreme climatic events did not have persistent effects on community structure or on soil nitrogen one year later. There was no consistent relationship between diversity and stability of biomass production of these plant communities, and species functional identity was the most effective explanation for the observed shifts in biomass production. The presence of C4 species resulted in an increased stability of productivity after extreme climatic events, but resulted in greater overall shifts in community composition. The presence of C4 species may buffer grassland community productivity against an increased frequency of extreme heating events associated with future global climate change.     

Short-term soil inorganic N pulse after experimental fire alters invasive and native annual plant production in a Mojave Desert shrubland

Post-fire changes in desert vegetation patterns are known, but the mechanisms are poorly understood. Theory suggests that pulse dynamics of resource availability confer advantages to invasive annual species, and that pulse timing can influence survival and competition among species. Precipitation patterns in the American Southwest are predicted to shift toward a drier climate, potentially altering post-fire resource availability and consequent vegetation dynamics. We quantified post-fire inorganic N dynamics and determined how annual plants respond to soil inorganic nitrogen variability following experimental fires in a Mojave Desert shrub community. Soil inorganic N, soil net N mineralization, and production of annual plants were measured beneath shrubs and in interspaces during 6 months following fire. Soil inorganic N pools in burned plots were up to 1 g m⁻² greater than unburned plots for several weeks and increased under shrubs (0.5–1.0 g m⁻²) more than interspaces (0.1–0.2 g m⁻²). Soil NO₃ ⁻−N (nitrate−N) increased more and persisted longer than soil NH₄ ⁺−N (ammonium−N). Laboratory incubations simulating low soil moisture conditions, and consistent with field moisture during the study, suggest that soil net ammonification and net nitrification were low and mostly unaffected by shrub canopy or burning. After late season rains, and where soil inorganic N pools were elevated after fire, productivity of the predominant invasive Schismus spp. increased and native annuals declined. Results suggest that increased N availability following wildfire can favor invasive annuals over natives. Whether the short-term success of invasive species following fire will direct long-term species composition changes remains to be seen, yet predicted changes in precipitation variability will likely interact with N cycling to affect invasive annual plant dominance following wildfire.     

Species evenness and invasion resistance of experimental grassland communities

Concern for biodiversity loss coupled with the accelerated rate of biological invasions has provoked much interest in assessing how native plant species diversity affects invasibility. Although experimental studies extensively document the effects of species richness on invader performance, the role of species evenness in such studies is rarely examined. Species evenness warrants more attention because the relative abundances of species can account for substantially more of the variance in plant community diversity and tend to change more rapidly and more frequently in response to disturbances than the absolute numbers of species. In this study, we experimentally manipulated species evenness within native prairie grassland mesocosms. We assessed how evenness affected primary productivity, light availability and the resistance of native communities to invasion. The primary productivity of native communities increased significantly with species evenness, and this increase in productivity was accompanied by significant decreases in light availability. However, evenness had no effect on native community resistance to invasion by three common exotic invasive species. In this study, niche complementarity provides a potential mechanism for the effects of evenness on productivity and light availability, but these effects apparently were not strong enough to alter the invasibility of the experimental communities. Our results suggest that species evenness enhances community productivity but provides no benefit to invasion resistance in otherwise functionally diverse communities.     

Two new Umbelliferae of the Chihuahuan Desert

Two apparently calcicolous species of apioid Umbelliferae,Aletes calcicola andDonnellsmithia coahuilensis, are described from the Chihuahuan Desert of Coahuila, Mexico.     

Functional redundancy in a clipping experiment on grassland plant communities

Functional redundancy predicts that some species may play equivalent roles in ecosystem functioning therefore conferring a kind of ‘insurance’ to perturbation when species richness is reduced, by the compensation of species of the same functional group on ecosystem processes. We evaluate functional redundancy on grassland plant communities by a removal experiment in which the evaluated treatments were: GG – clipping two graminoid species, FF – clipping two forb species, GF – clipping one graminoid and one forb species and Control – no removal. We tested the hypothesis that the above‐ground biomass removal of one species of each functional group would cause less change in the community composition (community persistence) and less decrease in biomass production than the above‐ground biomass removal of two species of the same functional group. Functional redundancy was corroborated for community persistence since treatments FG and C caused less change in community composition than treatments GG and FF, although no differences were found between treatments for above‐ground biomass. We verified that clipped species tend to be compensated by an increase in the percent cover of the remaining species of the same functional group. This work provides experimental evidence of early responses after plant clipping in small spatial scale of functional redundancy in naturally established grassland plant communities. We highlight redundancy as an intrinsic feature of communities insuring their reliability, as a consequence of species compensation within functional groups.     

Changes in plant species richness induce functional shifts in soil nematode communities in experimental grassland

Background

Changes in plant diversity may induce distinct changes in soil food web structure and accompanying soil feedbacks to plants. However, knowledge of the long-term consequences of plant community simplification for soil animal food webs and functioning is scarce. Nematodes, the most abundant and diverse soil Metazoa, represent the complexity of soil food webs as they comprise all major trophic groups and allow calculation of a number of functional indices.

Methodology/Principal Findings

We studied the functional composition of nematode communities three and five years after establishment of a grassland plant diversity experiment (Jena Experiment). In response to plant community simplification common nematode species disappeared and pronounced functional shifts in community structure occurred. The relevance of the fungal energy channel was higher in spring 2007 than in autumn 2005, particularly in species-rich plant assemblages. This resulted in a significant positive relationship between plant species richness and the ratio of fungal-to-bacterial feeders. Moreover, the density of predators increased significantly with plant diversity after five years, pointing to increased soil food web complexity in species-rich plant assemblages. Remarkably, in complex plant communities the nematode community shifted in favour of microbivores and predators, thereby reducing the relative abundance of plant feeders after five years.

Conclusions/Significance

The results suggest that species-poor plant assemblages may suffer from nematode communities detrimental to plants, whereas species-rich plant assemblages support a higher proportion of microbivorous nematodes stimulating nutrient cycling and hence plant performance; i.e. effects of nematodes on plants may switch from negative to positive. Overall, food web complexity is likely to decrease in response to plant community simplification and results of this study suggest that this results mainly from the loss of common species which likely alter plant – nematode interactions.     

Regional adaptation improves the performance of grassland plant communities

Many plant species are adapted locally or regionally. Whether such individual species performance translates into effects at community and ecosystem levels has rarely been tested. Such tests are crucial, however, to predict ecosystem consequences of sowing seed mixtures for grassland restoration or hay production. We compared the performance of replicated sown plant communities of regional origin with the performance of four foreign communities consisting of the same grassland species but originating from distances up to 890 km from our experimental site. The regional communities performed better than foreign communities in plant cover and diversity but not in aboveground biomass production. Additionally, in communities based on regional seeds fewer unsown species occurred and less bare ground was left open for erosion. Variation in community performance among source regions was related to climatic differences rather than to geographic distance to source regions. Individual species performance only partly explained community patterns, highlighting the importance of community level experiments. Our results suggest that the use of regional seeds represents an important approach to improve sown managed grasslands.     

Linking landscape history and dispersal traits in grassland plant communities

Dispersal limitation and long-term persistence are known to delay plant species’ responses to habitat fragmentation, but it is still unclear to what extent landscape history may explain the distribution of dispersal traits in present-day plant communities. We used quantitative data on long-distance seed dispersal potential by wind and grazing cattle (epi- and endozoochory), and on persistence (adult plant longevity and seed bank persistence) to quantify the linkages between dispersal and persistence traits in grassland plant communities and current and past landscape configurations. The long-distance dispersal potential of present-day communities was positively associated with the amounts of grassland in the historical (1835, 1938) landscape, and with a long continuity of grazing management—but was not associated with the properties of the current landscape. The study emphasises the role of history as a determinant of the dispersal potential of present-day grassland plant communities. The importance of long-distance dispersal processes has declined in the increasingly fragmented modern landscape, and long-term persistent species are expected to play a more dominant role in grassland communities in the future. However, even within highly fragmented landscapes, long-distance dispersed species may persist locally—delaying the repayment of the extinction debt.     

Effects of plant diversity on invasion of weed species in experimental pasture communities

Studies have shown that weed invasion into grasslands may be suppressed if the resident plant community is sufficiently diverse. The objective of this study was to determine whether increased forage plant diversity in grazed pasture communities might be associated with reduced weed abundance both in the aboveground vegetation and soil seed bank. Data were collected from a pasture experiment established in 1994 in Missouri, USA. The experiment consisted of 15 m×15 m plots sown with Festuca arundinacea Schreb. or Bromus inermis Leysser as a base species in mixtures of 1, 2, 3, 6, or 8 forage species. The plots were grazed by cattle during each growing season from 1998 to 2002. Aboveground plant species composition in each plot was measured using a point step method. Soil cores were collected in 1999 and 2002, and the species composition of germinable weed seeds in plots were evaluated by identifying seedlings as they germinated over an 8-week period. Species diversity was measured using several indices: species richness (S), Shannon–Wiener diversity index (H^′), and forage species evenness (J). Aboveground weed abundance in plots was unrelated to forage species richness (S), but weed abundance declined as the evenness (J) of resident forage species increased in mixtures. The species composition of mixtures may have affected weed abundance. Weeds both in the soil seed bank and aboveground vegetation were less abundant in mixtures that contained F. arundinacea compared with mixtures that contained B. inermis. Although variables like forage plant productivity may also suppress weed abundance in pastures, our results suggest that maintaining an evenly distributed mixture of forage species may help suppress weeds as well.

Zusammenfassung

Untersuchungen haben gezeigt, dass die Unkrautinvasion in Grünländer unterdrückt sein kann, wenn die ansässige Pflanzengemeinschaft ausreichend divers ist. Die Zielsetzung dieser Untersuchung war es zu bestimmen, ob eine erhöhte Futterpflanzendiversität in beweideten Grünlandgemeinschaften mit einer verringerten Unkrautabundanz sowohl bei der oberirdischen Vegetation als auch in der Bodensamenbank verbunden sein kann. Die Daten wurden in einem Weidelandexperiment gesammelt, das 1994 in Missouri, USA, etabliert wurde. Das Experiment bestand aus 15 m×15 m Probeflächen, die mit Festuca arundinacea Schreb. oder Bromus inermis Leysser als Basisarten in Mischungen von 1, 2, 3, 6 oder 8 Futterarten eingesät waren. Die Probeflächen wurden während jeder Wachstumssaison von 1998 bis 2002 stark mit Vieh beweidet. Die oberirdische Pflanzenartenzusammensetzung wurde in jeder Fläche mit einer Punktstopmethode gemessen. Bodenproben wurden 1999 und 2002 gesammelt und die Artenzusammensetzung der keimfähigen Unkrautsamen wurde in den Probeflächen bewertet, indem die Keimlinge identifiziert wurden, die in einer 8-wöchigen Periode keimten. Die Artendiversität wurde unter Verwendung verschiedener Indizes gemessen: Artenreichtum (S), Shannon–Wiener-Diversitätsindex (H^′) und Futterarten-Äquitabilität (J). Die oberirdische Unkrautartenabundanz in den Probeflächen stand in keiner Beziehung zum Futterartenreichtum (S), aber die Unkrautabundanz nahm ab, wenn die Äquitabilität (J) der ansässigen Futterarten in den Mischungen zunahm. Die Artenzusammensetzung der Mischungen könnte die Unkrautabundanz beeinflusst haben. Sowohl die Unkräuter in der Bodensamenbank, als auch in der oberirdischen Vegetation waren weniger abundant in Mischungen, die F. arundinacea enthielten, im Vergleich zu denen, die B. inermis enthielten. Obgleich Variablen wie die Futterpflanzenproduktivität möglicherweise ebenfalls die Unkrautabundanz im Weideland unterdrücken, lassen unsere Ergebnisse vermuten, dass die Aufrechterhaltung einer gleichmäßigen Mischung von Futterarten ebenfalls helfen kann, die Unkräuter zu unterdrücken.