Woody plant encroachment reduces species richness of herb‐rich woodlands in southern Australia

Abstract Woody plants have been increasing in many woodland and savanna ecosystems owing to land use changes in recent decades. We examined the effects of encroachment by the indigenous shrub Leptospermum scoparium (Myrtaceae) on herb‐rich Eucalyptus camaldulensis woodlands in southern Australia. Species richness and compositional patterns were examined under the canopy of L. scoparium and in surrounding open areas to determine the species most susceptible to structural changes. Richness was significantly lower in areas of moderate to high L. scoparium cover (>15%), suggesting that a threshold shrub cover caused major change in this ecosystem. Shrubs were associated with a significant reduction in above‐ground biomass of the ground‐layer flora and a significant shift in community composition. The few species that were positively associated with high L. scoparium cover were also common in the woodland flora; no new species were recorded under the shrub canopy. Important environmental changes associated with L. scoparium cover were decreased light availability and increased litter cover, which were likely a consequence of encroachment. Leptospermum scoparium cover was also associated with greater surface soil moisture, which may be a consequence of increased shading under the shrub canopy or indicate favourable soil conditions for L. scoparium establishment. Reductions in species richness and abundance of the germinable seed bank were found in soil samples taken from under L. scoparium. With ongoing recruitment of L. scoparium and consequent increases in shrub cover, ground‐layer diversity in these species‐rich woodlands should continue to decline over time.     

Drivers of understorey biomass: tree species identity is more important than richness in a young forest

林下生物量影响因素：幼龄林树种特性比丰富度更重要 生物多样性与生态系统功能的正相关关系已被广泛报道,其主要来源于对草原生态系统的研究。然而,该结论并不一定适用于更复杂的环境,例如具有不同垂直层次的森林。举例而言,已有研究表明上层乔木树种丰富度与林下生产力降低有关。树种丰富度是否会通过增加(由于生境异质性)或降低(通过增强竞争)资源的可利用性进而影响林下生产力,以及林下生产力是否受树种特性的影响更大,这些影响机制都可能会随着时间的推移而改变。此外,研究还表明,丰富度-生产力关系随着环境背景的变化而改变。本研究利用可以操控树种丰富度的实验林场研究了这些不同垂直层位里的时间和环境动态。在中国亚热带森林生物多样性与生态系统功能(BEF-China)研究计划的框架下,我们在3年时间里沿树种丰富度梯度反复采集林下生物量样本,研究了不同环境处理中树种丰富度、树种特性和时间对林下生物量的影响。尽管我们发现乔木层特性对林下生物量有显著和一致的影响,但是树种丰富度对后者却不具有这种影响。另外,在森林结构层之间,可能并不存在单一的、具有普遍性的上层乔木树种丰富度与林下生产力的相关关系,并且与上层乔木相关的环境因素(如透光率)对林下生产力的贡献程度会随着时间而变化。总体而言,我们的结果表明,在研究森林结构层之间的关系时应将时间动态变化考虑在内。     

Different inter‐annual responses to availability and form of nitrogen explain species coexistence in an alpine meadow community after release from grazing

Plant species and functional groups in nitrogen (N) limited communities may coexist through strong eco‐physiological niche differentiation, leading to idiosyncratic responses to multiple nutrition and disturbance regimes. Very little is known about how such responses depend on the availability of N in different chemical forms. Here we hypothesize that idiosyncratic year‐to‐year responses of plant functional groups to availability and form of nitrogen explain species coexistence in an alpine meadow community after release from grazing. We conducted a 6 year N addition experiment in an alpine meadow on the Tibetan Plateau released from grazing by livestock. The experimental design featured three N forms (ammonium, nitrate, and ammonium nitrate), crossed with three levels of N supply rates (0.375, 1.500 and 7.500 g N m^?2 yr^?1), with unfertilized treatments without and with light grazing as controls. All treatments showed increasing productivity and decreasing species richness after cessation of grazing and these responses were stronger at higher N rates. Although N forms did not affect aboveground biomas s at community level, different functional groups did show different responses to N chemical form and supply rate and these responses varied from year to year. In support of our hypothesis, these idiosyncratic responses seemed to enable a substantial diversity and biomass of sedges, forbs, and legumes to still coexist with the increasingly productive grasses in the absence of grazing, at least at low and intermediate N availability regimes. This study provides direct field‐based evidence in support of the hypothesis that idiosyncratic and annually varying responses to both N quantity and quality may be a key driver of community structure and species coexistence. This finding has important implications for the diversity and functioning of other ecosystems with spatial and temporal variation in available N quantity and quality as related to changing atmospheric N deposition, land‐use, and climate‐induced soil warming.     

From clear lakes to murky waters – tracing the functional response of high‐latitude lake communities to concurrent ‘greening’ and ‘browning’

Climate change and the intensification of land use practices are causing widespread eutrophication of subarctic lakes. The implications of this rapid change for lake ecosystem function remain poorly understood. To assess how freshwater communities respond to such profound changes in their habitat and resource availability, we conducted a space‐for‐time analysis of food‐web structure in 30 lakes situated across a temperature‐productivity gradient equivalent to the predicted future climate of subarctic Europe (temperature +3°C, precipitation +30% and nutrient +45 μg L^?1 total phosphorus). Along this gradient, we observed an increase in the assimilation of pelagic‐derived carbon from 25 to 75% throughout primary, secondary and tertiary consumers. This shift was overwhelmingly driven by the consumption of pelagic detritus by benthic primary consumers and was not accompanied by increased pelagic foraging by higher trophic level consumers. Our data also revealed a convergence of the carbon isotope ratios of pelagic and benthic food web endmembers in the warmest, most productive lakes indicating that the incorporation of terrestrial derived carbon into aquatic food webs increases as land use intensifies. These results, reflecting changes along a gradient characteristic of the predicted future environment throughout the subarctic, indicate that climate and land use driven eutrophication and browning are radically altering the function and fuelling of aquatic food webs in this biome.     

Large herbivores maintain a two‐phase herbaceous vegetation mosaic in a semi‐arid savanna

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Loss of Submerged Macrophytes in Shallow Lakes in North‐Eastern Germany

During the 1950s, the submerged vegetation of shallow lakes in north‐eastern Germany was dominated by nutrient tolerant species, with Ceratophyllum demersum and Myriophyllum sp. being most common. Almost one third of 300 investigated lakes had already lost their submerged macrophytes at that time. Very shallow lakes showed either high or low macrophyte abundance. Increasing depth resulted in medium macrophyte abundances, which may contribute to the stabilisation of local or temporary clearwater states. Forty years later, the percentage of lakes without macrophytes had dramatically increased. Between 55 and 85% of the investigated lakes showed a low abundance. The decline was most pronounced in very shallow lakes. The majority of the investigated lakes showed summer TP concentrations below 100 μg L^–1, but no colonisation by submerged macrophytes, which indicates a resilience against re‐colonisation.     

Cyanobacterial dynamics in shallow Lake Apopka (Florida,U.S.A.) before and after the shift from a macrophyte‐dominated to a phytoplankton‐dominated state

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Effects of smoke,heat and charred wood on the germination of dormant soil‐stored seeds from a Eucalyptus baxteri heathy‐woodland in Victoria,SE Australia

The effects of dry heat, wet heat, charred wood and smoke on the germination of dormant soil‐stored seeds from a Eucalyptus woodland in western Victoria were tested by using a glasshouse seed‐bank germination experiment. Seedling density, species richness and species composition were compared between replicated treated and control samples. A total of 5922 seedlings, comprising 59 plant species, was recorded from the soil samples over a period of 150 days. While a few species dominated (including Centrolepis strigosa, Wahlenbergia gracilenta and Ixodia achillaeoides), 26 species were represented by fewer than five seedlings and 18 species were restricted to single treatment types. With the exception of charred wood, all treatments led to a significant increase in seed germination relative to the control. The highest number of germinants was obtained for the smoke treatment, with a mean (± SE) of 12 547 ± 449 seedlings m^–2. Heat treatments yielded intermediate densities, with means (± SE) varying between 7445 ± 234 and 9133 ± 445 seedlings m^–2. In comparison with the estimates of seed‐bank sizes from other fire‐prone ecosystems, these densities are high. Species richness differed significantly among treatments. Highest mean richness was recorded in the smoke treatment and lowest for the control and charred wood treatments. There were significant differences in seed‐bank species composition between treatment types based on analysis of similarity (Anosim) using Bray–Curtis similarity. While heat was a specific requirement for triggering germination in hard‐seeded species (e.g. Fabaceae), smoke was the most effective trigger for species from a broad range of other families. The potentially confounding effect of physical and chemical mechanisms of germination stimulation in heated bulk soil samples is raised as an issue requiring further investigation in relation to the role of smoke as a germination trigger.     

Long‐term changes in the vegetation after the cessation of livestock grazing in Eucalyptus marginata (jarrah) woodland remnants

This paper documents changes in the floristic composition of Eucalyptus marginata Donn (jarrah) woodlands over 7 years of recovery from continual, intensive livestock grazing. In remnants of native woodland left after agricultural clearing, which have been subjected to livestock grazing, comparisons were made between the floristics of fenced exclosure plots and open plots that continued to be grazed. The vegetation in nearby remnants, which had not been subjected to livestock grazing, was also surveyed. An initial increase in annual exotic pasture species after grazing relief was only temporary and highly influenced by fluctuations in annual climatic patterns, particularly rainfall distribution and abundance. Subsequent years saw a decrease in exotic annuals in exclosure plots and an increase in native perennials, in a trend towards becoming more floristically similar to the ungrazed sites. Germination of overstorey species was observed in the exclosure plots, however, development of seedlings and saplings was sparse. Results indicate that for jarrah woodland in southwestern Australia, natural regeneration is possible after the removal of livestock, with the return (within 6 years) of native species richness to levels similar to those found in ungrazed vegetation. Re‐establishment of cover, however, appears to take longer. The floristic dynamics are described in terms of a nonequilibrium model. Two vegetation states exist, degraded remnants with an understorey dominated by annual species, and ungrazed vegetation with an understorey dominated by perennial shrubs and herbs. The former state is maintained by continual heavy grazing by livestock. Upon relief from grazing, the vegetation undergoes a transition towards floristic similarity to ungrazed vegetation. After 6 years, vegetation change in the exclosure plots appears to be continuing and therefore it is still in transition.     

Responses of woody vegetation to exclusion of large herbivores in semi‐arid savannas

The Nkuhlu large‐scale long‐term exclusion experiment in Kruger National Park was designed to study the long‐term effects of large herbivores on vegetation. One treatment excludes elephants, another excludes all herbivores larger than hares and another one comprises an open, control area. Vegetation monitoring was implemented in 2002 when a baseline survey was conducted prior to exclusion. Monitoring was repeated 5 years after exclusion. Data from the surveys were analysed to establish how structure and composition of woody vegetation had changed 5 years after herbivore exclusion. The analysis showed that neither plant assemblage nor mean vegetation height had changed significantly since exclusion. However, both species richness and density of woody plants increased 5 years after exclusion of all large herbivores, but not after the exclusion of elephants alone. One already common species, Dichrostachys cinerea, became more common after excluding all large herbivores compared with either no exclusion or elephant exclusion, possibly leading to competitive suppression of other species. Species other than D. cinerea tended to either increase or decrease in density, but the changes were insufficient to induce significant shifts in the overall assemblage of woody plants. The results indicate that after 5 years of exclusion, the combined assemblage of large herbivores, and not elephants alone, could induce changes in species richness and abundances of woody plants, but the effect was so far insufficient to induce measureable shifts in the assemblages of woody plants. It is possible that assemblages will change with time and increasing elephant numbers may amplify future changes.     

Near‐natural methods promote restoration of species‐rich grassland vegetation—revisiting a road verge trial after 9 years

The present loss of species‐rich grasslands makes it vital to restore these valuable habitat types, including novel habitat variants such as road verges. Due to the lack of knowledge on long‐term outcomes of restoration initiatives, well‐designed studies comparing different restoration methods are needed. In this study, we examined fine‐scale vegetation recovery patterns over 9 years in a field experiment with several near‐natural restoration methods (adding local seed mixtures, transferring hay from local grasslands using hard or light raking, and natural regeneration) in a road verge. We compared this to standard revegetation (hydroseeding species‐poor commercial seed mixtures). We found major temporal changes in vegetation restored by local seed or hay transfer, before it gradually became more similar to the donor grasslands and seed mixtures, which served as references for the experiment. Natural (spontaneous) regeneration with seed dispersal from surroundings gave similar results, whereas areas revegetated using standard methods became more dissimilar to the reference sites during the study period. The main variation in species composition reflected the contrast between local donor grasslands and seed mixtures and the species‐poor early successional grasslands. We conclude that near‐natural methods (hay transfer and seeding) successfully restored species‐rich grassland, including road verges. This study underlines the importance of comparing several treatments over a sufficiently long period to assess their success in restoring species‐rich grassland.     

Development of Brachystegia‐Julbernardia woodland after clear‐felling in central Zambia: Evidence for high resilience

Question: Does the development of Brachystegia‐Julbernardia (miombo) woodland after felling, and under a variable fire regime, occur via a serai stage of fire‐tolerant species? Location: Four sites in central Zambia, Africa. Methods: Trees in replicate plots were clear‐cut and stumps and resprouts enumerated. Species recruited into the tree layer (> 2.0 m tall) were monitored for 11 years (1991–2001) and fire occurrence and herbaceous biomass assessed annually to determine fuel loads. Results: Fire frequency was variable at the study sites and fuel loads were generally too low to suppress woodland regeneration after felling. However, at one site a change from low to high fire frequency arrested woodland development and triggered a regression towards a ‘fire‐trap’ vegetation type in which a few fire‐tolerant species survived. There was no evidence to support the hypothesis that miombo woodland regeneration is facilitated by a sere of fire‐tolerant species. All regrowth after felling was from resprouting plants present before felling. Trees with a previous history of felling sprouted more vigorously than trees that had not been felled before. Species richness in the tree layer increased with time since felling because resprout species had different height growth rates. Conclusion: The resilience of miombo trees after clear‐felling is largely due to their capacity to regenerate vegetatively from resprouts and stumps after release from frequent fires. Coppicing is therefore recommended as a suitable management technique for miombo woodland in central southern Africa.     

Estimating species richness and catch per unit effort from boat electro‐fishing in a lowland river in temperate Australia

Abstract Biodiversity estimates are typically a function of sampling effort and in this regard it is important to develop an understanding of taxon‐specific sampling requirements. Northern hemisphere studies have shown that estimates of riverine fish diversity are related to sampling effort, but such studies are lacking in the southern hemisphere. We used a dataset obtained from boat electro‐fishing the fish community along an essentially continuous 13‐km reach of the Murrumbidgee River, Australia, to investigate sampling effort effects on fish diversity estimates. This represents the first attempt to investigate relationships between sampling effort and the detection of fish species in a large lowland river in Australia. Seven species were recorded. Species‐specific patterns in catch per unit effort were evident and are discussed in terms of solitary and gregarious species, recreational fishing and the monitoring of rare and threatened species. There was a requirement to sample substantial lengths of river to describe total species richness of the fish community in this river reach. To this end, randomly allocated sampling effort and use of species richness estimators produced accurate estimates of species richness without the requirement for excessive levels of effort. Twenty operations were required to estimate species richness at this site, highlighting the need for comparable studies of river fish communities in lowland rivers elsewhere in Australia and the southern hemisphere.     

The impact of a fox‐ and cat‐free safe haven on the bird fauna of remnant vegetation in southwestern Australia

Introduced predators are a serious threat to Australian vertebrates. However, the consequences of predation for an area's avifauna have rarely been quantified. We took advantage of the establishment of a 7,832 ha fox‐ and cat‐free safe haven at Mt Gibson, in Western Australia, to assess the consequences of excluding introduced mammal predators on the bird fauna. Bird surveys were conducted over 6 years, before and after the establishment of the introduced predator‐free safe haven. After 3 years, half the sites were enclosed by the fence that excluded introduced predators, while the remainder of sites remained outside the fence and were exposed to fox and cat activity. The sites were stratified by four major vegetation types. A total of 91 bird species were variously detectable with the survey approach, but were typically more detectable during morning surveys. Site occupancy varied considerably among species, but overall, occupancy by all species was most likely to be either not impacted or positively impacted by the safe haven. The most notable change was that avifaunal richness appeared to increase in woodland and shrubland habitats within, as compared to outside, the safe haven. We conclude that: (1) the safe haven had an overall positive impact on bird occupancy; and (2) there were no consistent trends with respect to the kinds of species whose occupancy was positively impacted, beyond them all being small‐ to medium‐sized birds and mostly insectivorous. However, these conclusions must be tempered by the poor detection probability of many species.     

Non‐native species have multiple abundance–impact curves

The abundance–impact curve is helpful for understanding and managing the impacts of non‐native species. Abundance–impact curves can have a wide range of shapes (e.g., linear, threshold, sigmoid), each with its own implications for scientific understanding and management. Sometimes, the abundance–impact curve has been viewed as a property of the species, with a single curve for a species. I argue that the abundance–impact curve is determined jointly by a non‐native species and the ecosystem it invades, so that a species may have multiple abundance–impact curves. Models of the impacts of the invasive mussel Dreissena show how a single species can have multiple, noninterchangeable abundance–impact curves. To the extent that ecosystem characteristics determine the abundance–impact curve, abundance–impact curves based on horizontal designs (space‐for‐time substitution) may be misleading and should be used with great caution, it at all. It is important for scientists and managers to correctly specify the abundance–impact curve when considering the impacts of non‐native species. Diverting attention from the invading species to the invaded ecosystem, and especially to the interaction between species and ecosystem, could improve our understanding of how non‐native species affect ecosystems and reduce uncertainty around the effects of management of populations of non‐native species.     

Long‐term dynamics of submerged macrophytes and algae in a small and shallow,eutrophic lake: implications for the stability of macrophyte‐dominance

1. Submerged macrophyte and phytoplankton components of eutrophic, shallow lakes have frequently undergone dynamic changes in composition and abundance with important consequences for lake functioning and stability. However, because of a paucity of long‐term survey data, we know little regarding the nature, direction and sequencing of such changes over decadal–centennial or longer timescales. 2. To circumvent this problem, we analysed multiple (n = 5) chronologically correlated sediment cores for plant macro‐remains and a single core for pollen and diatoms from one small, shallow, English lake (Felbrigg Hall Lake, Norfolk, U.K.), documenting 250 years of change to macrophyte and algal communities. 3. All five cores showed broadly similar stratigraphic changes in macrophyte remains with three distinct phases of macrophyte development: Myriophyllum–Chara–Potamogeton (c. pre‐1900), to Ceratophyllum–Chara–Potamogeton (c. 1900–1960) and finally to Zannichellia–Potamogeton (c. post‐1960). Macrophyte species richness declined from at least 10 species pre‐1900 to just four species at the present day. Additionally, in the final Zannichellia–Potamogeton phase, a directional shift between epi‐benthic and phytoplankton‐based primary production was indicated by the diatom data. 4. Based on macrophyte–seasonality relationships established for the region, concomitant with the final shift to Zannichellia–Potamogeton, we infer a reduction in the seasonal duration of plant dominance (plant‐covered period). Furthermore, we hypothesise that this change in species composition resulted in a situation whereby macrophyte populations were seasonally ‘sandwiched’ between two phytoplankton peaks in spring and late summer as observed in the contemporary lake. 5. We suggest that eutrophication‐induced reductions in macrophyte species richness, especially if the number of plant‐seasonal strategies is reduced, may constrict the plant growing season. In turn, this may render a shallow lake increasingly vulnerable to seasonal invasions of phytoplankton resulting in further species losses in the plant community. Thus, as part of a slow (over perhaps 10–100s of years) and self‐perpetuating process, macrophytes may be gradually pushed out by phytoplankton without the need for a perturbation as required in the alternative stable states model of plant loss.     

Grazing Gradient versus Restoration Succession of Leymus chinensis (Trin.) Tzvel. Grassland in Inner Mongolia

Grazing‐induced degradation of grasslands is the primary impediment to the socioeconomic development of Inner Mongolia. It affects the entire environment of northern China. Understanding grassland dynamics is necessary for restoration and sustainable management of these degraded ecosystems. The recovery dynamics of a degraded Leymus chinensis (Trin.) Tzvel. grassland after removal of grazing was studied in comparison with its spatial variation along a grazing gradient, using its climax community as a benchmark. The species composition, diversity, and biomass of the grassland vegetation, as well as the attributes (height, density, and individual mass) of major species, were examined on the eight sites along the grazing gradient and in the recovering grassland over 11 years. The spatial pattern of grassland vegetation along the grazing gradient closely reflected its recovery trajectory over time. Both the spatial and the temporal processes exhibited the same shift in species dominance in association with grazing removal or less grazing intensity. Grassland degradation was accompanied by an increase in species density and a decrease in species size; this trend was reversed during recovery. This result suggested that the degraded grassland is highly resilient and that restoration could occur naturally by reducing or excluding grazing animals. However, some differences existed between the spatial and the temporal processes. Species richness was high on the light‐ or no‐grazing sites along the gradient, but varied little during the recovery of the degraded grassland. Species evenness was high under moderate to light grazing along the gradient and was high at the beginning of the recovery period but not at the end. Although standing biomass improved significantly during the recovery period, it did not change significantly along the grazing gradient. These observed discrepancies were related to the intrinsic difference in the spatial versus temporal processes and are discussed together with the advantage/disadvantage of the grazing gradient versus dynamic monitoring methods in grassland dynamics studies.