Changes in the vegetation of Quercus pubescens woodland after cessation of coppicing and grazing

Abstract. The cessation of coppicing and grazing in Quercus woodlands, with its subsequent changes in the structure, composition and functioning of vegetation communities, is becoming more frequent throughout the Mediterranean Basin. In southern France, we have studied successional changes in Quercus pubescens woodlands by visiting previously studied sites 18 yr later. Changes in vertical structure, species richness, floristic composition, life form and dispersal type were analysed and compared between woodlands that had previously been grazed or ungrazed. Both successions showed a decline in vegetation cover in the 0–25 cm height class and in the height class immediately under the canopy layer, due to oak litter accumulation and tree ageing. In post‐grazing succession, the abandonment of grazing and associated burning has allowed the vegetation cover to increase in the 0.25‐2 m height class. In both successions, grassland species decreased in frequency and forest species increased, a trend which was stronger in undisturbed succession. Species richness decreased with time in the undisturbed succession, but remained stable in the post‐grazing succession mainly because of the slow decline of plants linked to grazing. In undisturbed succession, therophytes and hemicryptophytes decreased. In contrast, therophytes and hemicryptophytes remained stable in post‐grazing succession. In both successions, endozoochorous species (notably Ruscus aculeatus) increased. Plants dispersed by non‐animal vectors decreased in undisturbed succession, but stability was observed in most of the dispersal types in post‐grazing succession. These results showed that a time‐lag existed between undisturbed succession and post‐grazing succession, the latter remaining at a younger stage of successional development due to more recent impact of grazing. However, both successions have converged suggesting that most of the traces of grazing on vegetation will disappear within a few years. The vegetation of these coppices, regardless of the previous grazing regime, will become increasingly similar to the vegetation of undisturbed woodlands. However, their floristic composition will probably never be identical to that of undisturbed woodlands, mainly because of the rarity of these undisturbed woodlands and of the short‐distance dispersal of many forest plant species.     

Long‐term vegetation stability and the concept of potential natural vegetation in the Neotropics

When vegetation trends over time are analysed from an appropriate long‐term perspective using palaeoecological records, the concept of potential natural vegetation (PNV) is unsupported because of continual vegetation changes driven by natural or anthropic forcings. However, some palaeoecological records show long‐lasting (i.e. millennial) vegetation stability at multidecadal to centennial time scales in the absence of natural and human drivers of change, which fits within the definition of PNV. A more detailed palaeoecological analysis of these records shows that they are an exception rather than a rule, and that they cannot be differentiated from other transient ecological states. Therefore, long records of vegetation stability cannot be considered to be valid evidence for PNV. From a palaeoecological perspective, the PNV concept is concluded to be unnecessary, even in cases of multidecadal to centennial vegetation stability in the absence of environmental disturbance.     

Long‐term changes in semi‐arid vegetation: Invasion of an exotic perennial grass has larger effects than rainfall variability

Abstract. Questions: This paper examines the long‐term change in the herbaceous layer of semi‐arid vegetation since grazing ceased. We asked whether (1) there were differences in the temporal trends of abundance among growth forms of plants; (2) season of rainfall affected the growth form response; (3) the presence of an invasive species influenced the abundance and species richness of native plants relative to non‐invaded plots, and (4) abundance of native plants and/or species richness was related to the time it took for an invasive species to invade a plot. Location: Alice Springs, Central Australia. Methods: Long‐term changes in the semi‐arid vegetation of Central Australia were measured over 28 years (1976–2004) to partition the effects of rainfall and an invasive perennial grass. The relative abundance (biomass) of all species was assessed 25 times in each of 24 plots (8 m × 1 m) across two sites that traversed floodplains and adjacent foot slopes. Photo‐points, starting in 1972, were also used to provide a broader overview of a landscape that had been intensively grazed by cattle and rabbits prior to the 1970s. Species’abundance data were amalgamated into growth forms to examine their relationship with environmental variation in space and time. Environmental variables included season and amount of rainfall, fire history, soil variability and the colonization of the plots by the exotic perennial grass Cenchrus ciliaris (Buffel grass). Results: Constrained ordination showed that season of rainfall and landscape variables relating to soil depth strongly influenced vegetation composition when Cenchrus was used as a covariate. When Cenchrus was included in constrained ordination, it was strongly related to the decline of all native growth forms over time. Univariate comparisons of non‐invaded vs impacted plots over time revealed unequivocal evidence that Cenchrus had caused the decline of all native growth form groups and species richness. They also revealed a contrasting response of native plants to season of rainfall, with a strong response of native grasses to summer rainfall and forbs to winter rainfall. In the presence of Cenchrus these responses were strongly attenuated. Discussion: Pronounced changes in the composition of vegetation were interpreted as a response to removal of grazing pressure, fluctuations in rainfall and, most importantly, invasion of an exotic grass. Declines in herbaceous species abundance and richness in the presence of Cenchrus appear to be directly related to competition for resources. Indirect effects may also be causing the declines of some woody species from changed fire regimes as a result of increased fuel loads. We predict that Cenchrus will begin to alter landscape level processes as a result of the direct and indirect effects of Cenchrus on the demography of native plants when there is a switch from resource limited (rainfall) establishment of native plants to seed limited recruitment.     

Arid vegetation in disequilibrium with livestock grazing: Evidence from long‐term exclosures

In recent decades, the conventional equilibrium paradigm for explaining rangeland vegetation dynamics has been challenged. Proponents of an alternative non‐equilibrium paradigm argue that in variable rangeland environments, external climatic events are critical to vegetation dynamics and there is little opportunity for plant–herbivore interactions to reach equilibrium. Understanding which paradigm more effectively describes an ecosystem has important consequences for management. In particular, some authors have argued that a focus on reducing stocking rates in non‐equilibrium systems may be futile, and management should be opportunistic in response to unpredictable rainfall events. We measured herbaceous biomass and plant species richness and abundance at five 14‐year exclosures on Innamincka Regional Reserve. Four were situated in the dunefields land system, and one on the Cooper Creek floodplain. We did not detect any significant differences between grazed and ungrazed treatments in total species richness or abundance, life form richness or abundance, or herbaceous biomass. Only one species, Portulaca oleracea, showed differences in abundance between treatments at more than one site, but the direction of these differences was not consistent. These results suggest that the non‐equilibrium paradigm more accurately describes vegetation dynamics in the dunefields and floodplains of north‐eastern South Australia. It is possible that some species had been lost from the study area prior to the establishment of the exclosures, thereby precluding recovery with protection from grazing; however, a regional analysis of the flora reveals little evidence of this. We argue that the dominance of ephemeral species confers resilience by limiting the development of strong feedbacks between grazing intensity and vegetation dynamics. Current grazing practices seem consistent with the conservation of plant species diversity across the dunefields and floodplains. Future studies should focus on the impacts of cattle grazing on areas of the landscape dominated by palatable perennials, as well as the small number of rare and potentially grazing‐sensitive species identified.     

The effect of long‐term repeated burning and fire exclusion on above‐ and below‐ground Blackbutt (Eucalyptus pilularis) forest vegetation assemblages

We used a long‐term fire experiment in south‐east Queensland, Australia, to determine the effects of frequent prescribed burning and fire exclusion on understorey vegetation (<7.5 m) richness and density in Eucalyptus pilularis forest. Our study provided a point in time assessment of the standing vegetation and soil‐stored vegetation at two experimental sites with treatments of biennial burning, quadrennial burning since 1971–1972 and no burning since 1969. Vegetation composition, density and richness of certain plant groups in the standing and soil‐stored vegetation were influenced by fire treatments. The density of resprouting plants <3 m in height was higher in the biennially burnt treatment than in the unburnt treatment, but resprouters 3–7.5 m in height were absent from the biennial burning treatment. Obligate seeder richness and density in the standing vegetation was not significantly influenced by the fire treatments, but richness of this plant group in the seed bank was higher in the quadrennial treatment at one site and in the long unburnt treatment at the other site. Long unburnt treatments had an understorey of rainforest species, while biennial burning at one site and quadrennial burning at the other site were associated with greater standing grass density relative to the unburnt treatment. This difference in vegetation composition due to fire regime potentially influences the flammability of the standing understorey vegetation. Significant interactions between fire regime and site, apparent in the standing and soil‐stored vegetation, demonstrate the high degree of natural variability in vegetation community responses to fire regimes.     

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.     

Edge effects across boundaries between natural and restored jarrah (Eucalyptus marginata) forests in south‐western Australia

Edge effects are a widespread and ubiquitous ecological phenomenon, yet they remain poorly studied across edges between restored and natural forests. To address this lack of knowledge, we studied vertebrate communities across edges between 3‐year old restored mine‐pits and adjacent unmined forest in the jarrah (Eucalyptus marginata) forest of south‐western Australia. We found that mammal communities showed no edge response but reptile communities did. Overall reptile abundance and Morethia obscura abundance were higher in unmined forest along edges, Egernia napoleonis abundance was lower in unmined forest along edges, while Pogona minor abundance was lower in restored mine‐pits along edges. Predictive models were unable to predict species edge responses, due to the lack of knowledge of the ecology of jarrah forest reptiles, but proved useful in identifying potential ecological mechanisms behind observed edge responses and suggested that potential mechanisms were likely different for each species. Our study is the first to show edge responses in both habitats forming the edge between restored and natural forests, emphasizing the importance of studying both habitats forming the edge. Our results also suggest that, despite being poorly studied, edge responses are common across edges between restored and natural forest and result from a variety of ecological mechanisms. An increased understanding of the ecological mechanisms driving edge responses across edges between restored and natural forests will improve our ability to integrate restored areas into cross‐landscape management and, ultimately, improve our ability to manage landscapes for biodiversity conservation.     

Recovery of understorey vegetation after release from a long history of sheep grazing in a herb‐rich woodland

The effects of stock grazing on native grassy ecosystems in temperate southern Australia are well documented. However, less is known about the potential of ecosystems to recover after a long history of stock grazing and, in particular, whether the removal of stock will have positive, negative or neutral impacts on biodiversity. We examined the response of understorey vegetation to the removal of sheep grazing in a herb‐rich Eucalyptus camaldulensis (red gum) woodland in western Victoria. Using a space‐for‐time chronosequence, woodlands were stratified into groups based on their time‐since‐grazing removal; these were long‐ungrazed (>20 years), intermediate‐time‐since‐grazing (9–14 years), recently ungrazed (5 years) and continuously grazed. We found significantly higher species density in long‐ungrazed sites relative to sites with a more recent grazing history. No differences were found in species density between continuously grazed sites and those ungrazed in the previous 14 years. Species composition differed with time‐since‐grazing removal and indicator species analysis detected several native species (including tall native geophytes and herbs) associated with long‐ungrazed sites that were absent or in low abundance in the more recently grazed sites. Seven of the eight species significantly associated with continuously grazed sites were exotic. Removal of sheep grazing in red gum woodlands can have positive benefits for understorey diversity but it is likely that recovery of key indicators such as native species will be slow.     

Long‐term productivity of Mediterranean herbaceous vegetation after a single phosphorus application

Question: What is the mechanism that underlies long‐term maintenance of high herbaceous productivity after a single application of phosphorus (4.5 gP m^?2 and 9 gP m^?2) in a hilly Mediterranean environment in a phosphorus‐deficient ecosystem? Location: Inland, 15 km E of the Mediterranean coast, W Galilee, Israel (35°15′E, 33°01′N; 500 m asl). Methods: The experiment was established in 1988. Multi‐year data on above‐ground biomass, botanical composition, P content of vegetation and soil, and the grazing management context of the experiment were integrated to construct a feasible account of the P dynamics of the ecosystem. Results: The productivity of the herbaceous component already responded to P application in the first year. The effect on the shrubby component of the ecosystem was marginal. The available (bicarbonate extractable) P in the upper soil layer peaked in the year after application of P and then declined to the original level within 7 years. Despite the decline in available soil P, a high, fluctuating level of herbaceous biomass production was maintained for 20 years. Legume species (Fabaceae) became a prominent constituent of the herbaceous vegetation after the P pulse. Conclusions: The long‐term shift in productivity of the herbaceous component of the grazed ecosystem was triggered by a nutritional pulse that induced a feedback loop based on changes in botanical composition of the herbaceous vegetation, the animal–vegetation interaction, grazing and supplementary feeding regimen of the cattle.     

Long‐term effects of the abandonment of grazing on steppe‐like grasslands

Question: What are the consequences of grazing abandonment on the Stipa lessingiana dominated steppe‐like grasslands? What is the relative importance of management and environmental factors in causing variation in species composition and abundance in the continuously grazed and abandoned grassland stands? Location: Transylvanian Lowland, Romania. Methods: Repeated vegetation mapping of a grassland stand, where grazing was abandoned 35 years ago; re‐sampling six grassland stands surveyed 29–57 years ago. For revealing long‐term changes in species composition and rank abundance PCoA ordination was applied. The relative importance of management and environmental factors in structuring vegetation were explored by CCA ordination. Diversity, evenness and the relative number and abundance of red‐listed species were compared between managed and abandoned stands. Results: Our results pointed out that grasslands which were formerly grazed and dominated by S. lessingiana, in the long‐term absence of grazing, have been transformed into a S. pulcherrima dominated type. Management, probably by creating bare surfaces and preventing litter accumulation, had the strongest effect on the species composition and abundance in the grasslands. Abandoned grassland stands had lower diversity and evenness compared to continuously grazed stands. While at the same time, the relative number of threatened, rare species did not differ between managed and abandoned sites. Conclusion: Maintaining extensively grazed, as well as un‐managed, Stipa dominated grasslands would be important in order to create various habitat conditions for plant species, especially threatened and rare species, and promote diversity on the landscape scale.     

Long‐term effects of exclusion of grazing stock on degraded herbaceous plant communities in a riparian Eucalyptus camaldulensis forest in south‐eastern Australia

Abstract Stock grazing has degraded many riparian ecosystems around the world. However, the potential for ecosystem recovery following the removal of grazing stock is poorly known. We developed a conceptual model to predict the responses of native and exotic herbaceous plants to grazing exclusion, based on site productivity and the degree of initial vegetation degradation. The effects of excluding grazing stock on richness, cover and composition of herbaceous plants were examined over 12 years in the degraded understorey of a riparian forest in Gulpa Island State Forest in south‐eastern Australia. We predicted that grazing exclusion would lead to limited changes in vegetation cover, richness and composition, owing to presumed low site productivity and the high degree of understorey degradation. Results showed that the cover, richness and composition of native and exotic species varied significantly among years. Over all plots, regions and years, total cover was slightly but significantly lower in grazed than in ungrazed plots (43.4% vs. 50.8%). While the cover of native plants increased over time in both treatments, the rate of increase was slightly greater in ungrazed plots. Grazing exclusion had no effect on the richness of native or exotic species, but had a significant but minor impact on plant composition, with different common species (mostly exotics) being promoted or diminished in ungrazed plots. The composition of grazed and ungrazed areas did not become more different over time. Overall, the effects that could be attributed to grazing exclusion were relatively minor and transient. Results are consistent with predictions based on site productivity and initial degradation, and should not be extrapolated to other more productive, or less degraded, riparian systems.     

Emergence of Eucalyptus marginata (jarrah) from seed in Mediterranean-climate forest in response to overstorey,site, seedbed and seed harvesting

Abstract The effects of overstorey, type of site, seedbed, seeding date and seed harvesting by vertebrates and invertebrates on the emergence of Eucalyptus marginata (jarrah) seedlings were studied experimentally in the northern jarrah forest of Western Australia. Seed harvesting by small vertebrates substantially reduced emergence, whereas invertebrates only slightly reduced emergence. Ninety per cent of seed was removed within 1 day when it was on the soil surface and in the open, whereas seed removal was insignificant when the seed was covered by soil. Vertebrates harvested seed more effectively when the understorey and litter were removed and the seed was more visible on the soil surface. Seed harvesting by small vertebrates had less impact on a site where there had been bauxite mining, possibly because the surrounding area was being revegetated with large numbers of seed and the area afforded poor cover for vertebrates. Emergence was less where the overstorey was removed compared to where it was retained, and on the rehabilitated bauxite mined site compared to the forest sites. There was no difference in emergence between the low and high quality forest sites and so the emergence phase could not fully explain the variation in abundance off. marginata seedlings on different quality sites. Post emergence events are likely to be important in explaining this difference between low and high quality sites.     

Effects of grazing and topography on long-term vegetation changes in a Mediterranean ecosystem in Israel

The dynamics of Mediterranean vegetation over 28 years was studied in the Northern Galilee Mountains, Israel, in order to identify and quantify the major factors affecting it at the landscape scale. Image analysis of historical and current aerial photographs was used to produce high resolution digital vegetation maps (pixel size = 30 cm) for an area of 4 km² in the Galilee Mountains, northern Israel. GIS tools were used to produce corresponding maps of grazing regime, topographic indices and other relevant environmental factors. The effects of those factors were quantified using a multiple regression analyses. Major changes in the vegetation occurred during the period studied (1964–1992); tree cover increased from 2% in 1964 to 41% in 1992, while herbaceous vegetation cover decreased from 56% in 1964 to 24% in 1992. Grazing, topography and initial vegetation cover were found to significantly affect present vegetation patterns. Both cattle grazing and goat grazing reduced the rate of increase in tree cover, yet even intensive grazing did not halt the process. Grazing affected also the woody-herbaceous vegetation dynamics, reducing the expansion of woody vegetation. Slope, aspect, and the interaction term between these two factors, significantly affected vegetation pattern. Altogether, 56% and 72% of the variability in herbaceous and tree cover, respectively, was explained by the regression models. This study indicates that spatially explicit Mediterranean vegetation dynamics can be predicted with fair accuracy using few biologically important environmental variables.     

Long‐term changes in ground beetle (Coleoptera: Carabidae) assemblages in Scotland

1. One‐way, directional changes in both plant and animal associations are likely to be occurring as a result of changing climate. Current knowledge of long‐term cycles in insect communities is scarce, and therefore it is difficult to assess whether the observed changes in insect communities are the first part of a long‐term trend or parts of normal cycles. 2. In this study multivariate methods were used to describe the trends in ground beetle (Coleoptera: Carabidae) assemblages over an 18‐year (1994–2011) period at two Scottish sites. In order to have a deeper insight into the underlying processes, both environmental factors and the species driving the detected changes were investigated. 3. In four out of the six sample transects, insect community compositions showed trends rather than fluctuating patterns. Hierarchical cluster analysis also revealed a clear separation, after accounting for sampling location and broad habitat, between early and later years of sampling. Decreasing annual maximum temperatures and increasing precipitation were identified as the main environmental drivers. Although increased rainfall was expected to be beneficial for hygrophilous species, in the transects in this study generalist species increased in dominance. 4. The increasing importance of generalists, in the communities studied here, underlines the vulnerability of the specialist species and urges greater effort in their conservation. Assemblage changes along different trajectories at the sites in the present study could only be tracked using multivariate methods; commonly used diversity indices proved to be unsatisfactory. Therefore, the exclusive use of simple diversity indices should be discouraged and multivariate methods should be preferred in environmental assessments and conservation planning.     

Long‐term body pigmentation changes on a manta ray (Mobulidae)

Manta rays (genus Manta) are listed as vulnerable on the IUCN Red List based partly on population studies that use body coloration and ventral spot patterns to identify and count individuals, as a result of such patterns being considered unique and permanent during their lifetime. The present study reports the first evidence of long‐term coloration changes of manta rays based on observations of one captive individual. Darkening skin pigmentation was documented on the side of the head, the inner side of the cephalic lobes and the pectoral fin margin of the ventral side, and spot markings appeared on the gill slits within 9 months. These changes were most likely ontogenetic or were connected to maturation. The described specimen's coloration did not fulfill the taxonomic classification criteria of either manta ray species and rather resembled two different manta ray species at the beginning and end of the study. These results show that coloration patterns of manta rays are not as stable as has been assumed previously. Further studies are needed to identify the extent of such changes for accurate identification and classification of manta rays. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 406–414.