Branch direction in Carnegiea gigantea (Cactaceae): Regional patterns and the effect of nurse plants

Abstract. Branch direction in Carnegiea gigantea (saguaro) cacti affects PAR interception and CO₂ uptake, and may influence reception of photomorphogenic cues. Branch direction data were collected in 29 saguaro populations over Arizona to determine if the southerly bias observed in previous studies at two locales can be generalized to the American range of the saguaro, and to determine the effect of shading by nurse plants on branch direction. Locally observed S bias from previous studies can be extended and generalized to include the American saguaro range. Branches were statistically uniform across environmental variability regarding temperature and precipitation. The S bias was not found in saguaros shaded by large nurse plants, supporting the assertion that branch direction occurs, in part, to maximize receipt of insolation.     

Effects of the Epichloë fungal endophyte symbiosis with Schedonorus pratensis on host grass invasiveness

Initial studies of grass–endophyte mutualisms using Schedonorus arundinaceus cultivar Kentucky‐31 infected with the vertically transmitted endophyte Epichloë coenophiala found strong, positive endophyte effects on host‐grass invasion success. However, more recent work using different cultivars of S. arundinaceus has cast doubt on the ubiquity of this effect, at least as it pertains to S. arundinaceus–E. coenophiala. We investigated the generality of previous work on vertically transmitted Epichloë‐associated grass invasiveness by studying a pair of very closely related species: S. pratensis and E. uncinata. Seven cultivars of S. pratensis and two cultivars of S. arundinaceus that were developed with high‐ or low‐endophyte infection rate were broadcast seeded into 2 × 2‐m plots in a tilled, old‐field grassland community in a completely randomized block design. Schedonorus abundance, endophyte infection rate, and co‐occurring vegetation were sampled 3, 4, 5, and 6 years after establishment, and the aboveground invertebrate community was sampled in S. pratensis plots 3 and 4 years after establishment. Endophyte infection did not enable the host grass to achieve high abundance in the plant community. Contrary to expectations, high‐endophyte S. pratensis increased plant richness relative to low‐endophyte cultivars. However, as expected, high‐endophyte S. pratensis marginally decreased invertebrate taxon richness. Endophyte effects on vegetation and invertebrate community composition were inconsistent among cultivars and were weaker than temporal effects. The effect of the grass–Epichloë symbiosis on diversity is not generalizable, but rather specific to species, cultivar, infection, and potentially site. Examining grass–endophyte systems using multiple cultivars and species replicated among sites will be important to determine the range of conditions in which endophyte associations benefit host grass performance and have subsequent effects on co‐occurring biotic communities.     

Interannual climate variability mediates changes in carbon and nitrogen pools caused by annual grass invasion in a semiarid shrubland

Exotic plant invasions alter ecosystem properties and threaten ecosystem functions globally. Interannual climate variability (ICV) influences both plant community composition (PCC) and soil properties, and interactions between ICV and PCC may influence nitrogen (N) and carbon (C) pools. We asked how ICV and non-native annual grass invasion covary to influence soil and plant N and C in a semiarid shrubland undergoing widespread ecosystem transformation due to invasions and altered fire regimes. We sampled four progressive stages of annual grass invasion at 20 sites across a large (25,000 km²) landscape for plant community composition, plant tissue N and C, and soil total N and C in 2013 and 2016, which followed 2 years of dry and wet conditions, respectively. Multivariate analyses and ANOVAs showed that in invasion stages where native shrub and perennial grass and forb communities were replaced by annual grass-dominated communities, the ecosystem lost more soil N and C in wet years. Path analysis showed that high water availability led to higher herbaceous cover in all invasion stages. In stages with native shrubs and perennial grasses, higher perennial grass cover was associated with increased soil C and N, while in annual-dominated stages, higher annual grass cover was associated with losses of soil C and N. Also, soil total C and C:N ratios were more homogeneous in annual-dominated invasion stages as indicated by within-site standard deviations. Loss of native shrubs and perennial grasses and forbs coupled with annual grass invasion may lead to long-term declines in soil N and C and hamper restoration efforts. Restoration strategies that use innovative techniques and novel species to address increasing temperatures and ICV and emphasize maintaining plant community structure—shrubs, grasses, and forbs—will allow sagebrush ecosystems to maintain C sequestration, soil fertility, and soil heterogeneity.     

Effects of invasion of fire‐free arid shrublands by a fire‐promoting invasive alien grass (Pennisetum setaceum) in South Africa

Arid shrublands in the Karoo (South Africa) seldom accumulate sufficient combustible fuel to support fire. However, as a result of invasion by an alien perennial grass (Pennisetum setaceum), they could become flammable. This paper reports on an experiment to assess the effects of fire following invasion by P. setaceum. We established 10 plots (5 × 10 m) separated by 2.5 m, and added grass fuel to five plots (5 and 10 tons ha^?1 to alternate halves of the plot) leaving the remaining five plots as interspersed controls. Plots with fuel added were burnt, and fire behaviour was measured during the burns. Rates of fire spread were generally low (0.01–0.07 m s^?1) and did not differ significantly between burn treatments. Mean fireline intensities were higher in the high compared with the low fuel treatments (894 and 427 kW m^?1, respectively). We recorded plant species and their cover before and after burning on each of the plots. After 15 months of follow‐up monitoring in the burn plots, only two species, the dwarf shrub (Tripteris sinuata) and the perennial herb (Gazania krebsiana) resprouted. Most individuals of other species were killed and did not reseed during the 15‐month study. The mass of added fuel load (high or low) did not influence vegetation recovery rates after fire. Should future invasions by P. setaceum lead to similar fuel loads in these shrublands, inevitable fires could change the vegetation and may favour spread of the flammable grass. Our results have important implications for predicting the effects of invasive alien plants (especially grasses) on fire‐free ecosystems elsewhere. The predicted impacts of fire may alter species composition, ultimately affecting core natural resources that support the Karoo economy.     

Invasion promotes invasion: Facilitation of C3 perennial grass dominance in mixed C3/C4 grassland by an invasive C3 woody sprouter (Prosopis glandulosa)

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Introduced annual grass increases regional fire activity across the arid western USA (1980–2009)

Non‐native, invasive grasses have been linked to altered grass‐fire cycles worldwide. Although a few studies have quantified resulting changes in fire activity at local scales, and many have speculated about larger scales, regional alterations to fire regimes remain poorly documented. We assessed the influence of large‐scale Bromus tectorum (hereafter cheatgrass) invasion on fire size, duration, spread rate, and interannual variability in comparison to other prominent land cover classes across the Great Basin, USA. We compared regional land cover maps to burned area measured using the Moderate Resolution Imaging Spectroradiometer (MODIS) for 2000–2009 and to fire extents recorded by the USGS registry of fires from 1980 to 2009. Cheatgrass dominates at least 6% of the central Great Basin (650 000 km²). MODIS records show that 13% of these cheatgrass‐dominated lands burned, resulting in a fire return interval of 78 years for any given location within cheatgrass. This proportion was more than double the amount burned across all other vegetation types (range: 0.5–6% burned). During the 1990s, this difference was even more extreme, with cheatgrass burning nearly four times more frequently than any native vegetation type (16% of cheatgrass burned compared to 1–5% of native vegetation). Cheatgrass was also disproportionately represented in the largest fires, comprising 24% of the land area of the 50 largest fires recorded by MODIS during the 2000s. Furthermore, multi‐date fires that burned across multiple vegetation types were significantly more likely to have started in cheatgrass. Finally, cheatgrass fires showed a strong interannual response to wet years, a trend only weakly observed in native vegetation types. These results demonstrate that cheatgrass invasion has substantially altered the regional fire regime. Although this result has been suspected by managers for decades, this study is the first to document recent cheatgrass‐driven fire regimes at a regional scale.     

Restoration of C4 grasses with seasonal fires in a C3/C4 grassland invaded by Prosopis glandulosa,a fire‐resistant shrub

Questions: Can prescribed fire restore C₄ perennial grasses in grassland ecosystems that have become dominated by fire‐resistant C₃ shrubs (Prosopis glandulosa) and C₃ grasses? Do fires in different seasons alter the direction of change in grass composition? Location: Texas, USA. Methods: We quantified short‐ and long‐term (12 yr post‐fire) herbaceous functional group cover and diversity responses to replicated seasonal fire treatments: (1) repeated‐winter fires (three in 5 yr), (2) repeated‐summer fires (two in 3 yr), and (3) alternate‐season fires (two winter and one summer in 4 yr), compared with a no‐fire control. Results: Summer fires were more intense than winter fires, but all fire treatments temporarily decreased Prosopis and C₃ annual grass cover. The alternate‐season fire treatment caused a long‐term increase in C₄ mid‐grass cover and functional group diversity. The repeated‐summer fire treatment increased C₄ short‐grass cover but also caused a long‐term increase in bare ground. The repeated winter fire treatment had no long‐term effects on perennial grass cover. Mesquite post‐fire regrowth had increasingly negative impacts on herbaceous cover in all fire treatments. Conclusions: Summer fire was necessary to shift herbaceous composition toward C₄ mid‐grasses. However, the repeated‐summer fire treatment may have been too extreme and caused post‐fire herbaceous composition to “over‐shift” toward less productive C₄ short‐grasses rather than C₄ mid‐grasses. This study provides some of the first long‐term data showing a possible benefit of mixing seasonal fires (i.e., the alternate‐season fire treatment) in a prescribed burning management plan to restore C₄ mid‐grass cover and enhance overall herbaceous diversity.     

Plant–plant interactions as a mechanism structuring plant diversity in a Mediterranean semi‐arid ecosystem

Plant–plant interactions are among the fundamental processes that shape structure and functioning of arid and semi‐arid plant communities. Despite the large amount of studies that have assessed the relationship between plant–plant interactions (i.e., facilitation and competition) and diversity, often researchers forget a third kind of interaction, known as allelopathy. We examined the effect of plant–plant interactions of three dominant species: the perennial grass Lygeum spartum, the allelopathic dwarf shrub Artemisia herba‐alba, and the nurse shrub Salsola vermiculata, on plant diversity and species composition in a semi‐arid ecosystem in NE Spain. Specifically, we quantified the interaction outcome (IO) based on species co‐occurrence, we analyzed diversity by calculation of the individual species–area relationship (ISAR), and compositional changes by calculation of the Chao‐Jaccard similarity index. We found that S. vermiculata had more positive IO values than L. spartum, and A. herba‐alba had values between them. Lygeum spartum and A. herba‐alba acted as diversity repellers, whereas S. vermiculata acted as a diversity accumulator. As aridity increased, A. herba‐alba transitioned from diversity repeller to neutral and S. vermiculata transitioned from neutral to diversity accumulator, while L. spartum remained as diversity repeller. Artemisia herba‐alba had more perennial grass species in its local neighborhood than expected by the null model, suggesting some tolerance of this group to its “chemical neighbor”. Consequently, species that coexist with A. herba‐alba were very similar among different A. herba‐alba individuals. Our findings highlight the role of the nurse shrub S. vermiculata as ecosystem engineer, creating and maintaining patches of diversity, as well as the complex mechanism that an allelopathic plant may have on diversity and species assemblage. Further research is needed to determine the relative importance of allelopathy and competition in the overall interference of allelopathic plants.     

Sampling grain influences trends in vegetation composition and diversity with time since fire in Australian heathland

Scale‐dependency of pattern and process is well‐understood for many ecological communities; however, the influence of spatial scale (sampling grain) in detecting temporal change in communities is less well‐understood. The temperate lowland heathlands of south‐east Australia are one of the most fire‐prone ecosystems on earth. Despite the extensive literature documenting the effect of time since fire on heathlands, we know little about how sampling grain influences trends in vegetation variables over time, and whether these trends are scale‐dependent. Using 3500 ha of heathland in the Gippsland Lakes Coastal Park, south‐east Australia, we investigated how above‐ground species composition and diversity, and trends in these variables with increasing time since fire, were influenced by sampling grain (1 m², 10 m², 100 m², 900 m², 1 ha, 4 ha). Sampling grain influenced patterns detected in vegetation variables and in some instances, significantly affected their relationship with time since fire. Richness decreased with time since fire, with mean richness decreasing at three of the four grains, while total richness decreased at half of the sampled grains. Evenness (J) decreased with increasing time since fire for all grains except 1 m². The decline in diversity (H) with time since fire appeared to be independent of scale, as all grains decreased significantly with increasing time since fire. Community heterogeneity demonstrated a weak response to time since fire across most grains. Changes in composition among young (0–6 years since fire), intermediate (9–19 years) and old (23–27 years) sites were dependent on sampling grain, with all grains exhibiting significant differences in composition, apart from the 1 m² grain and the 100 m² grain (presence/absence data). Overall, species composition, richness, evenness, diversity and community heterogeneity were dependent on the scale at which the vegetation was sampled. In addition, trends in many of these vegetation variables with increasing time since fire were scale‐dependent. This work provides strong evidence that sampling at multiple grains contributes substantially to understanding pattern and process in heathlands.     

Decomposition and nutrient release of grass and tree fine roots along an environmental gradient in southern Patagonia

Decomposition of fine roots is a fundamental ecosystem process that relates to carbon (C) and nutrient cycling in terrestrial ecosystems. However, this important ecosystem process has been hardly studied in Patagonian ecosystems. The aim of this work was to study root decomposition and nutrient release from fine roots of grasses and trees (Nothofagus antarctica) across a range of Patagonian ecosystems that included steppe, primary forest and silvopastoral forests. After 2.2 years of decomposition in the field all roots retained 70–90% of their original mass, and decomposition rates were 0.09 and 0.15 year^?1 for grass roots in steppe and primary forest, respectively. For N. antarctica roots, no significant differences were found in rates of decay between primary and silvopastoral forests (k = 0.07 year^?1). Possibly low temperatures of these southern sites restricted decomposition by microorganisms. Nutrient release differed between sites and root types. Across all ecosystem categories, nitrogen (N) retention in decomposing biomass followed the order: tree roots > roots of forest grasses > roots of steppe grasses. Phosphorus (P) was retained in grass roots in forest plots but was released during decomposition of tree and steppe grass roots. Calcium (Ca) dynamics also was different between root types, since trees showed retention during the initial phase, whereas grass roots showed a slow and consistent Ca release during decomposition. Potassium (K) was the only nutrient that was rapidly released from both grass and tree roots in both grasslands and woodlands. We found that silvopastoral use of N. antarctica forests does not affect grass or tree root decomposition and/or nutrient release, since no significant differences were found for any nutrient according to ecosystem type. Information about tree and grass root decomposition found in this work could be useful to understand C and nutrient cycling in these southern ecosystems, which are characterized by extreme climatic conditions.     

Plant community recovery after herbicide management to remove Phragmites australis in Great Lakes coastal wetlands

Invasive plants, such as Phragmites australis, are a global threat to plant diversity and are commonly controlled using herbicide management. The purpose of this study was to evaluate the plant community response 6–10 years after large‐scale herbicide management to remove Phragmites from Great Lakes coastal wetlands along the shores of western Lake Erie. Vegetation surveys were conducted in nine wetlands undergoing herbicide management and four unmanaged Phragmites‐dominated wetlands. The relative percent cover of Phragmites was dramatically lower in the managed (1.3%) compared to unmanaged wetlands (93.0%; p < 0.001), although relative percent cover of other non‐natives following herbicide management averaged 39.2% (ranging from 6.4 to 67.6%). The cover‐weighted floristic quality index was significantly higher in managed wetlands (p < 0.01), with the highest indices (12.4–17.0) at sites that received prescribed fire after herbicide treatment (p < 0.05). Species richness and diversity were significantly higher in managed wetlands (p < 0.001); however, there was no significant difference between wetlands treated only with herbicide and those treated with herbicide and prescribed fire. Our results indicate that herbicide management is effective in reducing Phragmites and improving floristic quality over timescales of 6–10 years. However, continued spot‐treatment and management of new invasive species may be required, and the return of high‐quality plant communities may be unrealistic in the study region.     

To burn or not to burn: Comparing reintroducing fire with cutting an encroaching conifer for conservation of an imperiled shrub‐steppe

Woody vegetation has increased on rangelands worldwide for the past 100–200 years, often because of reduced fire frequency. However, there is a general aversion to reintroducing fire, and therefore, fire surrogates are often used in its place to reverse woody plant encroachment. Determining the conservation effectiveness of reintroducing fire compared with fire surrogates over different time scales is needed to improve conservation efforts. We evaluated the conservation effectiveness of reintroducing fire with a fire surrogate (cutting) applied over the last ~30 years to control juniper (Juniperus occidentalis Hook.) encroachment on 77 sagebrush‐steppe sites. Critical to conservation of this imperiled ecosystem is to limit juniper, not encourage exotic annual grasses, and promote sagebrush dominance of the overstory. Reintroducing fire was more effective than cutting at reducing juniper abundance and extending the period of time that juniper was not dominating the plant community. Sagebrush was reduced more with burning than cutting. Sagebrush, however, was predicted to be a substantial component of the overstory longer in burned than cut areas because of more effective juniper control. Variation in exotic annual grass cover was explained by environmental variables and perennial grass abundance, but not treatment, with annual grasses being problematic on hotter and drier sites with less perennial grass. This suggests that ecological memory varies along an environmental gradient. Reintroducing fire was more effective than cutting at conserving sagebrush‐steppe encroached by juniper over extended time frames; however, cutting was more effective for short‐term conservation. This suggests fire and fire surrogates both have critical roles in conservation of imperiled ecosystems.     

Effects of an African grass invasion on Hawaiian shrubland nitrogen biogeochemistry

African perennial C₄ grasses are highly successful invaders in Hawaiian ecosystems. We examined the effects of African molasses grass (Melinis minutiflora Beauv.) on Hawaiian shrubland nitrogen (N) dynamics without the influence of fire disturbance. Vegetation tissue carbon and nitrogen chemistry, soil inorganic N pools, net N mineralization rates, and total soil N were studied in three adjacent areas: a monospecificMelinis grassland, a mixed grass/shrubland mosaic, and an un-invaded shrubland.Melinis plots within the mosaic area exhibited the largest inorganic N pools and fastest net N mineralization rates, but were temporally variable with grass phenology. Un-invaded shrubland plots contained the smallest inorganic N pools and lowest net N mineralization rates. Grass foliar C:N and litter C:N were lower than those of common shrubland species, providing one possible link between species and ecosystem N dynamics at this site. The combined effects of N cycle modification, successful light competition, and fire-cycle enhancement make the invasion ofMelinis a significant perturbation to Hawaiian shrubland ecosystem function and successional dynamics. ei]Section editor H Lambers     

Use of saguaro fruit by white-winged doves: isotopic evidence of a tight ecological association

We report the use of stable isotope and crop content analyses to quantify the use of saguaro (Carnegiea gigantea) nectar and fruit by migratory desert white-winged doves (Zenaida asiatica mearsnii). Saguaro resources had characteristically ¹³C-enriched CAM values (δ¹³C=–12.8±0.7‰ SD VPDB and –13.1±0.5‰ SD VPDB for nectar and fruit, respectively) relative to other food plants used by doves (δ¹³C_C3=–24.9±3.3‰ SD VPDB). The water contained in saguaro nectar and fruit was deuterium enriched (δD=19.6±2.0‰ SD VSMOW and 48.4±1.6‰ SD VSMOW for nectar and fruit, respectively) relative to other water sources (ranging from –41 to –19‰ VSMOW). During the fruiting season, there was a positive correlation between δ¹³C in dove liver tissues and percent of saguaro in crop contents. A two-point mixing model indicated that during the peak of saguaro fruit use, most of the carbon incorporated in dove tissues was from saguaro. Desert white-winged doves appear to be saguaro specialists. Averaged over the period when doves were resident, saguaro comprised about 60% of the total carbon incorporated into dove tissues. Tissue δ¹³C and δD of body water showed a significant positive correlation, indicating that doves were using saguaro as a source of both nutrients and water. However, at the peak of saguaro utilization, the doves’ body-water δD was more positive (by about 20‰) than saguaro fruit water. We hypothesize that this enrichment is due to fractionated evaporative water losses by doves. Using dove carbon isotope data and a two end-point mixing model we estimate that, on average, doves consume the equivalent of 128 saguaro fruits per season; each fruit contains on average 26.0±14.8 g SD of pulp (wet mass) of which 19.4 g is water. Stable isotopes have been used to produce qualitative re-constructions of animal diets. Our study shows that they can be used to provide quantitative estimates of the flow of nutrients from resources into consumers as well. Received: 30 September 1999 / Accepted: 23 March 2000     

Insect assemblages change along a gradient of invasion by a nonnative grass

Because invasions by nonnative plants alter the structure and composition of native plant communities, invasions can alter the function of ecosystems for animals that depend on plants for food and habitat. We quantified effects of an invasion by a nonnative grass on the insect community in grasslands of southeastern Arizona. We sampled insects on 54 1-ha plots established across a gradient of invasion by Lehmann lovegrass (Eragrostis lehmanniana Nees), a perennial species native to southern Africa. Between 2000 and 2004, we captured 94,209 insects representing 13 orders, 91 families, and 698 morphospecies during 2,997 trap nights. Richness of families, richness of morphospecies, and overall abundance of insects decreased as dominance of nonnative grass increased. With every 100 g/m² increase in biomass of nonnative grass, the average number of insect families decreased by 5%, morphospecies decreased by 6%, and overall abundance decreased by 14%. In areas dominated by nonnative grass, 2 of 8 orders and 6 of 27 families of insects were present less frequently and one family was present more frequently; 5 of 8 orders and 6 of 27 families of insects were less abundant and 3 families were more abundant than in areas dominated by native grasses. As a result, this plant invasion altered the structure of the insect community, which has consequences for animals at higher trophic levels and for ecosystem processes, including decomposition and pollination. Because complete eradication of nonnative plants might be possible only rarely, maintaining stands of native vegetation in invaded areas may be an important practical strategy to foster persistence of animals in grasslands invaded by nonnative plants.     

Dominant Grasses Suppress Local Diversity in Restored Tallgrass Prairie

Warm‐season (C₄) grasses commonly dominate tallgrass prairie restorations, often at the expense of subordinate grasses and forbs that contribute most to diversity in this ecosystem. To assess whether the cover and abundance of dominant grass species constrain plant diversity, we removed 0, 50, or 100% of tillers of two dominant species (Andropogon gerardii or Panicum virgatum) in a 7‐year‐old prairie restoration. Removing 100% of the most abundant species, A. gerardii, significantly increased light availability, forb productivity, forb cover, species richness, species evenness, and species diversity. Removal of a less abundant but very common species, P. virgatum, did not significantly affect resource availability or the local plant community. We observed no effect of removal treatments on critical belowground resources, including inorganic soil N or soil moisture. Species richness was inversely correlated with total grass productivity and percent grass cover and positively correlated with light availability at the soil surface. These relationships suggest that differential species richness among removal treatments resulted from treatment induced differences in aboveground resources rather than the belowground resources. Selective removal of the dominant species A. gerardii provided an opportunity for seeded forb species to become established leading to an increase in species richness and diversity. Therefore, management practices that target reductions in cover or biomass of the dominant species may enhance diversity in established and grass‐dominated mesic grassland restorations.     

Fire and torrential rainfall: effects on the perennial grass Brachypodium retusum

Summer wildfires and autumn torrential rainfall are the background to the development of one of the most important environmental problems in the western Mediterranean area: erosion, degradation of soil structure and desertification. These processes especially affect plant communities located in old abandoned fields where woody sprouters are practically absent. Under these conditions, post-fire vegetation cover is dependent on obligate seeders and, in the short term, especially on resprouting perennial grasses. Brachypodium retusum is a rhizomatous perennial grass that plays a particularly important role in the resilience of these fire-prone Mediterranean ecosystems. In Mediterranean gorse shrublands, during the first few years after a fire, this species represents 90% of plant cover and biomass. The recurrence or severity of fire does not seem to affect the regeneration capacity of this species. After two years, even under high fire frequency (12 years) and high fire severity (> 400°C), it recovers both cover and biomass. Nevertheless, this species, a key factor in the control of soil erosion, is extremely affected by erosion processes. A single extreme rainfall event reduces the rate of regeneration of B. retusum by about 50%. The reduction in the protective cover of this species may result in degradation and eventual self-induced degrading process in the ecosystem.     

Prefire grazing by cattle increases postfire resistance to exotic annual grass (Bromus tectorum) invasion and dominance for decades

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