Interaction of livestock grazing and rainfall manipulation enhances herbaceous species diversity and aboveground biomass in a humid savanna

Journal of Plant Research - Understanding of the interaction of livestock grazing and rainfall variability may aid in predicting the patterns of herbaceous species diversity and biomass production....     

Influence of rainfall and grazing on herbage dynamics in a seasonally dry tropical savanna

Species composition and herbage dynamics in relation to rainfall variability and cattle grazing were studied in permanently protected, grazed, and temporarily fenced treatments on three sites in a seasonally dry tropical savanna. Permanently protected sites, established between 1979 and 1984, were 55–79% similar with each other in species composition, and 14–25% similar with grazed sites during the period 1986–1988. Similarity among grazed sites was only 36–43%. Number of species was greater in the grazed treatment than in the permanently protected treatment. The percentages of annual grasses and non-leguminous forbs were greater in grazed savanna than in permanently protected savanna. Species diversity was higher in grazed savanna than in the corresponding permanently protected savanna. Species the two annual cycles studied, peak live shoot biomass was 614 g m^-2 in permanently protected savanna, 109 g m^-2 in grazed savanna, and 724 g m^-2 in temporarily fenced savanna. Live shoot biomass in temporarily fenced savanna was 18 to 44% greater than in permanently protected savanna. Peak canopy biomass ranged from 342 to 700 g m^-2 in permanently protected savanna. It was related with total rainy season rainfall, and was particularly sensitive to late rainy season rainfall. On the other hand, peak canopy biomass in grazed savanna ranged from 59 to 169 g m^-2 and was related to grazing intensity rather than either total rainy season rainfall or late rainy season rainfall. Coefficient of variation of green biomass in permanently protected savanna was related with rainfall variability indicating it to be a pulsed system which responds quickly to rainfall events. Biomass of woody species ranged from 2466 to 5298 g m^-2 in permanently protected savanna and from 744 to 1433 g m^–2 in the grazed savanna. Green foliage biomass was 3.7 to 6.4% of the woody biomass in permanently protected and 5.6 to 5.9% in grazed savanna, and supplements substantially the fodder resource during the dry periods of the year.     

Influence of rainfall amount and livestock grazing on soil respiration in a moist Kenyan savannah

The influence of livestock grazing and changing rainfall patterns on soil respiration (Rs) in the tropical savannah in Lambwe is unclear. Measurements of Rs using chambers were conducted between June and November 2012 in a humid savannah in Western Kenya to determine how simulated rainfall amounts and cattle grazing affected soil respiration. The experimental layout was split plot comprising rainfall simulations which were laid on grazed and fenced blocks, respectively. The rainfall simulations split plots each measured 6 m × 3 m and were covered with rainout gutters above the canopy of the herbaceous layer to either increase or reduce ambient rainfall by 50%. Grazing increased soil bulk density and significantly (p < 0.05) lowered soil respiration rates and soil water content irrespective of the rainfall treatment. The positive correlation between Rs and root biomass in both grazed and fenced plots revealed a significant contribution of root growth activities to Rs. Increased soil water content (SWC) by rainfall addition improved water penetration into deeper soil layers and therefore stimulated root and microbial activities which in turn built up soil respiration. Reduction in SWC by rainfall exclusion lowered microbial contact with the organic substrate and likely caused death and/ or inactivity of roots.     

Effect of low rainfall and browsing by large herbivores on an enclosed savannah habitat in Kenya

Savannah ecosystems in East Africa are rarely stable and can experience rapid local changes from dense woodlands to open plains. In this 3‐year study there was a reduction of 16.3% in a height‐stratified sample of nearly 1000 individually marked Acacia drepanolobium trees. The study was carried out in an enclosed fire‐free wooded grassland habitat in the Laikipia region of Kenya. The trees were monitored from 1998 to 2001, a period that included 12 months when rainfall was 60% below average. Elephants were responsible for the loss of 40% of the trees, black rhinos 33% and 27% died from the effects of the drought. Low rainfall was correlated with increased damage as elephants switched diet from grass to trees. Heavy browsing by giraffes reduced tree growth rates and increased their susceptibility to drought. Hence the combination of low rainfall and heavy browsing by elephants, black rhinos and giraffes led to the rapid tree loss. These findings have implications for research into the causes of instability in savannah ecosystems and the management of enclosed reserves.     

Scattered trees and livestock grazing as keystones organisms for sustainable use and conservation of Mediterranean dehesas

Mediterranean scattered oak woodlands support relatively high biological diversity, and provide important ecosystem services. However, there is still a lack of knowledge about livestock-tree-grassland relationships, knowledge necessary for effective conservation management in these systems. To address such lacuna, we measured the effects of scattered trees and grazing animals (livestock vs. wild ungulates, mostly deer) on biomass and diversity (and their relationship) of the herbaceous layer (understory) of open holm oak woodland (known as dehesa) in Central Spain, for two years. We located two 1-ha plots within three different grazing management schemes (cattle, sheep and wildlife only). Within each plot, we randomly selected four focal trees from which a subplot was established in two directions and three distances. In each subplot, we measured plant diversity and biomass, along with microclimatic variables across sites. We found rainfall variability affected herbaceous biomass and diversity in the dehesa system, and the effects were nuanced: in a dry year (<300 mm annual rainfall) the presence of livestock grazing correlated with higher herbaceous biomass (14%), and in moister year (>600 mm annual rainfall) the effect was enhanced, as plots under livestock grazing exhibited even higher herbaceous biomass (42%). In addition, livestock, particularly cattle, generated a more diverse community (52 species m⁻²). Microsites created by trees generated high plant diversity among herbaceous communities, where alpha diversity was higher (20%) in ecotone and inter-tree gaps than under the canopies. In addition, species turnover was higher than 50% in all cases. Biomass was higher (15%) under the canopies only under humid conditions. Legume dry matter was positively related to plant diversity. We concluded that agroforestry management practices such as promoting tree development through silvicultural techniques to create different microsites, and maintaining livestock grazing are needed to preserve these emblematic ecosystems.     

Exclusion of livestock grazing and wood collection in dryland savannah: an effect on long‐term vegetation succession

Sahelian savannah faces increasing pressure from human activities, leading to its degradation. The aim of this study was to investigate the possibility of restoration of dryland savannah vegetation by the elimination of disturbance factors on the ecosystem. Is degraded dryland savannah vegetation able to be restored by means of natural succession? What is the timescale for its recovery? The study took place in the Bandia Reserve, 65 km south‐east of Dakar (Senegal), a unique site with two successional stages due to the elimination of uncontrolled exploitation. The vegetation structure of 15 years (15YRS) and 5 years (5YRS) after fencing was compared with vegetation exposed to continuous livestock grazing and wood collection outside the fenced area. Calculated by redundancy analysis, a significant effect of selected areas on the cover of all species was revealed and successional stage explained more than 45% of data variability. Perennial forbs, annual forbs and perennial grasses achieved the highest cover in 5YRS, woody species in 15YRS and annual grasses in the area outside of the fenced reserve. The dominant woody species Acacia seyal, A. ataxacantha, A. nilotica subsp. adstringens and Balanites aegyptiaca reconstituted the dense formation of Acacia bushland by means of natural succession in the 15YRS area.     

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.     

天坑森林植物群落叶功能性状、物种多样性和功能多样性特征

以天坑内部-边缘-外部森林植物群落为研究对象,通过调查植物的群落结构、叶功能性状,探究天坑内外森林植物群落叶功能性状、物种多样性和功能多样性变化特征及其内在关联,为深入了解负地形森林生态系统的功能和恢复退化喀斯特地区的植被提供一定参考。研究结果如下：（1）比叶面积（SLA: 198.75 cm2/g)）、叶面积（LA: 42.70 cm2）、叶磷含量（LPC: 1.70 g/kg）和叶钾含量（LKC: 10.27 g/kg）在天坑内部最高,叶组织密度（LTD: 0.32 g/cm3）和叶干物质含量（LDMC: 0.41 g/g）在天坑外部最高,天坑内外森林均易受到磷限制,表明随天坑内部-边缘-外部生境变化,植物对环境的适应机制和生存策略发生了部分调整,物种的防御策略增强,生长投入策略减弱。（2）Shannon-Wiener指数（2.82）、Simpson指数（0.92）和Pielou’s均匀度指数（0.87）均以天坑外部最高,功能丰富度（1.05）、功能离散度（1.88）和Rao’s二次熵（4.52）以天坑内部最高,表明随天坑内部-边缘-外部生境的变化,植物功能性状的差异减少,物种分布及其功能性状分布总体上更为均匀、物种数量增多。（3）物种多样性指数之间、功能多样性指数之间存在较强的相关性,表明物种多样性指数之间、功能多样性指数之间存在不同的制约关系。（4）叶功能性状与物种多样性、功能多样性的相关性强,物种多样性和功能多样性之间相关性较弱,表明叶性状对生态学过程的变化较为敏感,叶功能性状与物种多样性之间存在较强的耦合关系。     

Spatial heterogeneity in the herbaceous layer of a semi-arid savanna ecosystem

Despite increasing recognition of the role spatial pattern can play in ecosystem function, few studies have quantified spatial heterogeneity in savanna ecosystems. The spatial distribution of herbaceous biomass and species composition was measured across three scales in a semi-arid savanna in central Kenya, and patterns were related to environmental variables at different scales. Herbaceous biomass declined across a rainfall gradient and from upper to lower topographic positions, but variation within a site (across 5–50 m) was similar in magnitude to among-site variation associated with rainfall and topography. Geostatistical analyses showed that patchiness at scales of 5–25 m explained 20% of total variation in herbaceous biomass. This pattern arose from the presence of both 5–10-m diameter patches containing high herbaceous biomass (> 170 g m^–2) and 5–10-m diameter patches characterized by nearly bare soil surfaces (< 40 g m^–2). Patch structure was contingent on topography, with larger bare patches at ridgeline and upper hillslope positions. Grass species distributions showed the greatest degree of patch structure and species turnover across distances of 5–45 m. Additional community variation was associated with topography, with minimal variation in species composition across the rainfall gradient. Pattern diversity significantly exceeded levels reported for four other grassland ecosystems, suggesting fundamental differences in local processes generating spatial pattern. It is hypothesized that heterogeneously distributed grazing pressure, interacting with the distribution of shrub canopies, is an important factor generating such high levels of small-scale patch structure in this savanna.     

Effects of grazing exclusion on plant species richness and phytomass accumulation vary across a regional productivity gradient

Question: Does long‐term grazing exclusion affect plant species diversity? And does this effect vary with long‐term phytomass accumulation across a regional productivity gradient? Location: Lowland grassy ecosystems across the state of Victoria, southeast Australia. Methods: Floristic surveys and phytomass sampling were conducted across a broad‐scale productivity gradient in grazing exclusion plots and adjacent grazed areas. Differences in species richness, evenness and life‐form evenness between grazed and ungrazed areas were analysed. The environmental drivers of long‐term phytomass accumulation were assessed using multiple linear regression analysis. Results: Species richness declined in the absence of grazing only at the high productivity sites (i.e. when phytomass accumulation was >500 g m^?2). Species evenness and life‐form evenness also showed a negative relationship with increasing phytomass accumulation. Phytomass accumulation was positively associated with both soil nitrogen and rainfall, and negatively associated with tree cover. Conclusions: Competitive dominance is a key factor regulating plant diversity in productive grassy ecosystems, but canopy disturbance is not likely to be necessary to maintain diversity in less productive systems. The results support the predictions of models of the effects of grazing on plant diversity, such as the dynamic equilibrium model, whereby the effects of herbivory are context‐dependent and vary according to gradients of rainfall, soil fertility and tree cover.     

Dry grassland plant diversity conservation using low-intensity sheep and goat grazing management: case study in Prague (Czech Republic)

After abandonment, dry grassland (Festuco-Brometea) areas decline due to gradual overgrowing by woody species and the expansion of perennial tall grass species. Dry grassland vegetation was formed by extensive livestock grazing, thus grazing is considered one of the most natural methods for managing this type of vegetation. Six years after introducing low-intensity sheep and goat grazing in seven nature reserves in Prague (Czech Republic), the following impact of this management on dry grassland vegetation was observed: The cover of expansive woody species, particularly Ligustrum vulgare, and to a smaller extent Cornus sanguinea and Prunus spinosa declined. In addition, a significant, long-term declining trend of the expansive species Arrhenatherum elatius was also observed. Also the cover of Pimpinella saxifraga and Allium senescens declined significantly with regard to statistical evaluation. On the contrary, the cover of Achillea millefolium, Centaurea stoebe, Securigera varia, Elytrigia repens, Erysimum crepidifolium, Falcaria vulgaris, Fallopia convolvulus and Verbascum lychnitis increased. The cover of species characteristic of dry grasslands (Festuco-Brometea) increased significantly. No changes were observed in the number and cover of the Red List species. In addition, the presence of nitrophilous and ruderal species increased. Species diversity also significantly increased. From our findings we can conclude that managing dry grasslands with low-intensity grazing can help to keep dry grassland vegetation in good condition and conserve its plant diversity. Nomenclature: Kubát et al. (2002) for taxa and Moravec et al. (1995) for syntaxa.     

The drivers of woody species richness and density in a Neotropical savannah

Environmental filtering prevents species without certain attributes from occurring in local communities. Traits respond differently to different abiotic factors, assembling communities with varying composition along environmental gradients. Here, we measured proxies of soil fertility, disturbance by fire, response and physiological traits to assess how these variables interact to determine woody species richness and density in a Neotropical savannah. We explicitly incorporated our assumptions about how different abiotic filters influence different subsets of traits into a statistical model using structural equation modelling, yielding a more accurate representation of the assembly process. Fire had an effect on resistance traits, whereas soil fertility influenced physiological traits. Resistance traits explained both the richness and density of plots, whereas physiological traits explained only the density. Fewer fire events led to richer and denser plots. Similarly, areas with lower cation exchange capacity assembled less dense communities. Furthermore, we showed that structural equation modelling yielded a realistic representation of the bivariate interactions of distinct environmental filters with different subsets of traits.     

Conflicting management objectives on the Colorado Plateau: Understanding the effects of bison and cattle grazing on plant community composition

The American Bison (Bison bison Linnaeus) in the Henry Mountains are one of the last free-roaming, genetically pure herds of bison remaining in North America. Anecdotal evidence indicates that this herd is utilising a cattle winter range during the summer and fall, creating a conflict between the state agency that manages the bison, and the Bureau of Land Management (BLM) and local ranchers. In theory, the addition of bison grazing pressure could reduce forage availability in the short term and lead to undesired changes in the plant community in the long term. Our objective was to determine whether bison have altered the plant species composition of the cattle winter range. We characterised plant species composition, percent cover, and grazing intensity on three adjacent, geomorphologically similar mesas. Grazing regimes were different on the three mesas, one with bison and cattle present, one with cattle only present, and the third with neither cattle nor bison present. Vegetation surveys were accompanied by a 28-year remote sensing time series to test for temporal shifts in an index of primary productivity. We found a higher grazing intensity on two dominant forage species on the bison plus cattle grazed mesa in fall, before the cattle were turned out to winter pasture. Despite this difference in grazing intensity, we found few differences in species composition, percent cover, or NDVI across the three grazing regimes. Our results suggest that high intensity summer bison grazing, while likely creating short-term reductions in forage availability, has not caused differences in plant community composition or productive potential. Shifts in community composition can take years to unfold and just as long to correct; therefore, continued monitoring of the combined effects of cattle and bison is needed.     

植物群落和物种多样性动态过程及物种丧失机制放牧试验

在长期严格的沙地草场放牧试验的基础上,通过调查及多种计算和分析,揭示了在放牧条件下植物群落和物种多样性的动态演变规律及物种丧失的机制,主要结论是:(1)在不同稳定的牧压下群落外貌特征及组分发生明显的分异。(2)持续强度放牧使物种消失,α多样性下降,为理解现代物种消亡和灭绝机制提供了部分依据,现代物种的消亡和灭绝正是由于人们对于生境的破坏引起的,生境破坏引起局部地区物种的消失,这种局部消失的不断累加就可能引起物种的消亡和绝灭。(3)放牧试验对物种组成和β多样性的影响表现在时空两个方面。     

Effects of open grazing and livestock exclusion on floristic composition and diversity in natural ecosystem of Western Saudi Arabia

Livestock grazing is one of the main causes of rangeland degradation in Saudi Arabia. Fencing to exclude grazers is one of the main management practices used to restore vegetation and conserve biodiversity. The main objectives of this study were to investigate the changes in plant diversity and abundance, floristic composition and plant groups of the major life forms in response to thirty-five years of grazing exclosure in western Saudi Arabia. These vegetation attributes and palatability were compared in 30 sampling stands located in the excluded and grazed sites. Our results showed that livestock exclusion significantly increased covers, density and species richness of annuals, grasses, perennial forbs, shrubs and trees. Exclosure enhanced the abundance and richness of palatable species and depressed the development of weedy species. About 66.7% of the recorded species at the excluded site were highly palatable compared to 34.5% at the grazed site. In contrary, about 55.2% unpalatable species were found in the grazed site compared to 25.8% in the protected site. Jaccard’s similarity index between the excluded and grazed sites showed lower values of 0.39%, 0.40% and 0.31% at levels of families, genus and species, respectively. The results suggest that establishing livestock exclusion may be a useful sustainable management tool for vegetation restoration and conservation of plant diversity in degraded rangelands of arid regions.     

Carbon dioxide exchange and biomass productivity of the herbaceous layer of a managed tropical humid savanna ecosystem in western Kenya

Aims Humid savannas, as a result of high precipitation amounts, are highly productive. They are also hotspots for land use change and potential sources of carbon dioxide (CO₂) due to the large soil carbon (C) stocks. Understanding how ecosystem CO₂ exchange is influenced by changes arising from agricultural land use is vital in future management of these ecosystems and in responding to the ongoing shifts in management and climate. The aim of this study was to identify how ecosystem CO₂ exchange and biomass productivity of the herbaceous layer of a humid savanna in Kenya respond to current management practices.Methods We used flux chambers to quantify CO₂ fluxes, while monthly harvests were undertaken to determine biomass development of the herbaceous layer of three sites that were (i) fenced to exclude livestock grazing, (ii) subjected to grazing by livestock and (iii) abandoned after being cultivated for maize production and also open to grazing by livestock.Important findings The peak aboveground biomass ranged between 380 and 1449g m ?2 and biomass production was significantly (P < 0.05) lower in the grazed and abandoned plots. The maximum gross primary production (GPP) and net ecosystem CO₂ exchange (NEE) ranged between 21.8±1.3 to 32.5±2.7 and ?9.6±0.7 to^-17.9±4.8 μmol m ?2 s^-1, respectively. Seasonal NEE fluctuations ranged between 10 and 21 μmol m ?2 s^-1, while spatial (among sites) differences ranged between 2 and 10 μmol m ?2 s^-1. Ecosystem respiration (R eco) fluctuated between 5 and 10 μmol m ?2 s^-1 during the growing season. R eco was, however, not significantly different among the sites. Unlike in other similar ecosystems where ecosystem respiration is determined by the ambient temperature, we did not find any relationship between R eco and temperature in this savanna. Instead, soil moisture accounted for 38–88% of the spatial and seasonal fluctuations in ecosystem CO₂ fluxes and aboveground biomass production. Management influenced the maximum GPP and NEE rates through modification of soil moisture, plant species composition and aboveground biomass. We concluded that soil moisture is the key determinant of ecosystem CO₂ exchange and productivity in this tropical savanna. Management, however, significantly modifies C fluxes and productivity through its influence on soil moisture, plant species composition and aboveground green biomass and should be taken into consideration in future similar studies.