Predicting invasion dynamics of four alien Pinus species in a highly fragmented semi-arid shrubland in South Africa

This study explored the determinants of spread of four alien Pinus species and the ability of models to predict invasion dynamics in a complex fragmented landscape. The role of environmental factors, natural and anthropogenic disturbance in relation to invasion history was assessed for different stages in the invasion process using a Geographic Information System. Pines escaped from plantations over the past 30 years and spread into the natural semi-arid shrubland (renosterveld). The pattern of spread was compared with a simulated random distribution using two different techniques, a standard logistic regression, and a new recursive modelling approach (Formal Inference-based Recursive Modelling; FIRM). FIRM analysis improved the accuracy of predictions and revealed interactive effects of variables hidden by the logistic regression analysis. More than 80% of isolated pine individuals were found in 20% of the habitat classified as suitable by the models. Soil pH was the most important predictor for the distribution of isolated trees, whereas the establishment of dense pine stands was largely determined by fire history. Differences in invasive behaviour could be explained by species attributes such as limited dispersal for P. canariensis, and better drought-tolerance for P. halepensis. Sixty-five percent of the current pine distribution was accurately predicted by the spatial distribution of the first trees to have invaded. Such models could be used to predict potential spread of invasive plants and gain a better understanding of the main factors driving the invasion process. However, the spread of invasive species in fragmented landscapes, strongly modified by human activities, is very complicated, and the spread remains difficult to predict in the long term. The dynamics of invasion are discussed in relation to changes in land use and disturbance regime.     

Effects of experimental warming and drought on biomass accumulation in a Mediterranean shrubland

We studied the effects of experimental warming and drought on the plant biomass of a Mediterranean shrubland. We monitored growth at plant level and biomass accumulation at stand level. The experimentation period stretched over 7 years (1999–2005) and we focused on the two dominant shrub species, Erica multiflora L. and Globularia alypum L. and the tree species Pinus halepensis L. The warming treatment increased shoot elongation in E. multiflora, and the drought treatment reduced shoot elongation in G. alypum. The elongation of P. halepensis remained unaffected under both treatments. The balance between the patterns observed in biomass accumulation for the three studied species in the drought plots (reduction in E. multiflora and P. halepensis and increase in G. alypum) resulted in a trend to reduce 33% the biomass of the drought treatment plots with respect to the untreated plots, which almost doubled their biomass from 1998 to 2005. The results also suggest that under drier conditions larger accumulation of dead biomass may occur at stand level, which combined with higher temperatures, may thus increase fire risk in the Mediterranean area.     

Response of mangroves to drought and non-tidal conditions in St Lucia Estuary,South Africa

The effect of a prolonged closed-mouth state on the condition of mangrove habitats was studied in May 2010 at St Lucia Estuary, KwaZulu-Natal. The mouth had been closed to the sea since 2002 as a result of artificial mouth closure and drought, which had resulted in non-tidal, dry and hypersaline aquatic conditions. Sediment characteristics, mangrove population structure, and snail and crab abundance, were measured at four sites of differing salinity and water level to determine if there was a relationship between environmental characteristics and biotic response. A site fringing the main channel had the lowest elevation and the highest density of mangrove seedlings and saplings at the water's edge, whereas a dry site at the highest elevation had the lowest density of mangroves with mostly tall adult trees and few saplings. At these two sites the salinity (>34.5) was significantly higher and the surface sediment water content (<30%) was significantly lower than at other sites. Highest tree density was found at a freshwater seepage site, where all size classes were represented, while the permanently flooded site between the Mfolozi River and St Lucia Estuary had the tallest trees but no seedlings or saplings. At the freshwater seepage and permanently flooded sites, porewater and sediment salinity (<10) were significantly lower, while sediment moisture (>55.5%) was significantly higher, than at the fringing and dry sites. Low sediment moisture was unfavourable for mangrove growth and recruitment. Long-term monitoring is needed to document the response of the mangroves to closed-mouth, non-tidal conditions.     

Local responses to conservation in the Richtersveld National Park, South Africa

Local notions of conservation deserve closer scrutiny. We cannot assume that they naturally coincide with those of the tourist core; nor that they are automatically given higher priority when local populations are induced to participate in practices which conserve flora and fauna. A case study of the establishment of the Richtersveld National Park in Namaqualand, a remote arid region of South Africa, is used to argue that local attitudes to conservation and development (associated with tourism) can become increasingly cynical and ambivalent in the context of ecotourism. Furthermore, such attitudes are shaped less by economic inducements than by the often contradictory messages about conservation that are being communicated daily.     

Contrasting growth changes in two dominant species of a Mediterranean shrubland submitted to experimental drought and warming

BACKGROUND AND AIMS: Climate projections predict drier and warmer conditions in the Mediterranean basin in the next decades. The possibility of such climatic changes modifying the growth of two Mediterranean species, Erica multiflora and Globularia alypum, which are common components of Mediterranean shrublands, was assessed. METHODS: A field experiment was performed from March 1999 to March 2002 to prolong the drought period and to increase the night-time temperature in a Mediterranean shrubland, where E. multiflora and G. alypum are the dominant species. Annual growth in stem diameter and length of both species was measured and annual stem biomass production was estimated for 1999, 2000 and 2001. Plant seasonal growth was also assessed. KEY RESULTS: On average, drought treatment reduced soil moisture 22 %, and warming increased temperature by 0.7-1.6 degrees C. Erica multiflora plants in the drought treatment showed a 46 % lower annual stem elongation than controls. The decrease in water availability also reduced by 31 % the annual stem diameter increment and by 43 % the annual stem elongation of G. alypum plants. New shoot growth of G. alypum was also strongly reduced. Allometrically estimated biomass production was decreased by drought in both species. Warming treatment produced contrasting effects on the growth patterns of these species. Warmer conditions increased, on average, the stem basal diameter growth of E. multiflora plants by 35 %, raising also their estimated stem biomass production. On the contrary, plants of G. alypum in the warming treatment showed a 14 % lower annual stem growth in basal diameter and shorter new shoots in spring compared with controls. CONCLUSIONS: The results indicate changes in the annual productivity of these Mediterranean shrubs under near future drier and warmer conditions. They also point to alterations in their competitive abilities, which could lead to changes in the species composition of these ecosystems in the long term.     

Disturbance,drought and dynamics of desert dune grassland,South Africa

A seven-year study of marked plants and plots in Stipagrostis ciliata (Desf.) de Winter dune grassland, in the arid (<100 mm yr^–1) Bushmanland area of the Northern Cape province of South Africa, was designed to test the hypothesis that establishment of ephemeral plants, and recruitment of perennial grasses was dependent upon disturbances that reduced the density of living perennial grass tussocks. In 1989, eight 4 m² plots were cleared of perennial vegetation by uprooting and removing all plants so as to resemble small-scale disturbances made by burrowing mammals or territorial antelope. The vegetation on the cleared plots and surroundings was monitored until 1996. Initial results supported our hypothesis. In wet years, when ephemeral plants were abundant, their average fresh mass was 2–3 times greater per unit area on the cleared plots than in control plots in adjacent, undisturbed grassland. Many Stipagrostis seedlings established in the cleared plots over the two years following clearing but were rare in adjacent areas among established conspecifics. However, a drought in 1992 (11 mm of rain over 12 months) lead to widespread mortality of the perennial grass, killing 56% (range 22–79%) of established tufts. High densities of Stipagrostis seedlings appeared following the drought-breaking rains in January 1993, both in the disturbed plots and in the surrounding `undisturbed' dune grassland. Ephemeral plants established in large numbers throughout the area during the high rainfall year of 1996 and were generally more numerous in the old disturbances than in control plots. Seven years after clearing the biomass of grass on the cleared plots was approximately 34% of the mass removed from the plots in 1989 whereas in the undisturbed grassland biomass was 66% of 1989 levels. Drought had little long-term effect on community composition, and Stipagrostis ciliata constituted 94–98% of plant community before and after drought. Cleared plots were recolonised by S. ciliata, but the contribution of other grass species increased by 6–9%. Synchronous recruitment following occasional drought-induced mortality can generate even-aged populations of the dominant desert dune grasses.     

Experimental drought and warming decrease diversity and slow down post-fire succession in a Mediterranean shrubland

Plant community recovery (species richness, diversity and composition) of a post-fire Mediterranean shrubland was monitored over a seven year period (1998–2005) under experimental drought and warming that simulated the environmental conditions forecast for this area in the coming decades. Species richness and Shannon's index were positively correlated with accumulated precipitation in the growing season and both variables were negatively affected by reduced water availability in drought plots. The relative abundance of the different species in both treatments was linearly correlated with their relative abundance in control plots. Moreover, we found species-specific responses to treatments. Drought and warming treatment reduced the competitive ability of the obligate seeder tree Pinus halepensis against native resprouter shrubs and consequently, the transformation from shrub to pine tree dominated vegetation was slowed down. Conversely, the water use strategy of Globularia alypum may allow this species to maintain a dominant position in drought plots. Therefore, future drier and warmer conditions in Mediterranean areas may severely affect plant community recovery after a disturbance, due to the existence of both abundance-dependent and species-specific responses that may change inter-specific competitive relationships.     

Warming and drought alter soil phosphatase activity and soil P availability in a Mediterranean shrubland

We conducted a field experiment simulating the warming and drought in a Mediterranean shrubland dominated by Erica multiflora and Globularia alypum with the aim to simulate the next future climate conditions predicted by the IPCC and ecophysiological models. As P is frequently a limiting nutrient in Mediterranean ecosystems, we investigated the drought and warming effects on soil phosphatases activities, soil P contents and availability, litter and leaf P concentration, and the capacity of this community to maintain soil P reserves and retain this nutrient in the ecosystem. Warming treatment increased soil and air temperature (an average of 1°C) and drought treatment decreased soil water content in one of the seasons analysed (28% in autum 2004). Warming increased (68%) the activities of soil acid phosphatases in summer and alkaline phosphatase activity (22%) in spring 2004, and increased P concentrations in E. multiflora. Instead, warming decreased P concentrations in litterfall of this same species, E. multiflora, and soil HCO₃-extractable P_i (Olsen-P_i) in some seasons, decreasing total P soil concentration (37%) after 6 years of treatment. The drought treatment did not change soil phosphatase activities, nor available P_i. The effects of climate change on soil P dynamics in Mediterranean areas will thus be strongly dependent on whether the main variable involved in the local change is warming or drought. If warming is the main change without significant changes in water availability, the increases of biological activity can accelerate plant growth, P capture by plants and increase soil-phosphatase activity, altogether decreasing P contents in soil. If drought is the main change, a reduction in P demands by plants is expected, increasing P stocks in soils.     

Disturbances in semi-arid shrubland and arid grassland in the Karoo, South Africa: mammal diggings as germination sites

Mammals (mainly aardvark Orycteropus ufer , porcupine Hystrix austro-africanae , bat-eared fox Otocyon megalotis and Cape fox Vulpes chama ) which excavate holes in hard-capped soils in Karoo shrubland and inter-dune pans create microsites where seeds, detritus and water accumulate, and seeds germinate. Holes with seedlings were significantly deeper (6.7 ± 0.4 cm) than holes without seedlings (4.5 ± 0.3 cm). In open, inter-shrub sites, significantly more seedlings germinated in diggings than on the flat soil surface, but there was no difference in the proportion of seedlings germinating under shrubs and in diggings. In inter-dune pans, 96% of all seedlings found were growing in diggings.

Résumé

Les mammifères (surtout l'oryctérope Orycreropus afer , le porcépic Hystrix austro-ajricanae , l'otocyon Otocyion megalotis , et le renard du Cap Vulpes chama ) qui creusent des trous dans les régions buissonneuses du Karoo et dans les creux entre les dunes créent des microsites où les semences, les déchets et l'eau se rassemblent, et les semences germent. Les trous où elles germent sont significativement plus profonds (6,7 ± 0,4 cm) que ceux où elles ne germent pas (4,5± 0,3 cm). Aux endroits buissonneux ouverts, il pousse plus de jeunes plantes dans des creux qu'à la surface du sol, mais il n'y a pas de différence entre les proportions de celles qui poussent sous les buissons et dans les creux. Dans les creux entre les dunes, 96% des jeunes pousses croissent dans les excavations.     

The density and stability of birds in shrubland and drainage line woodland in the southern Karoo,South Africa

Comparative surveys of bird species richness and abundance showed that a total of 69 bird species was recorded on 1 km transects (n = 996) through shrubland, with a total estimated density of birds of 32.7 ± 32.8 (S.D.) birds/km², of which resident birds made up 24.4 ± 21.0 birds/km², nomads made up 1.9 ± 11.7 birds/km² and local nomads made up 6.3 ± 20.1 birds/km². There was no correlation in general between total bird numbers or total bird biomass and rain, but numbers and biomass of nomads showed the strongest correlations with rainfall two months previously. A total of 86 species was recorded on 276 surveys along a 1 km stretch of drainage line woodland. Species richness varied from an average of about nine species to 24 species each month, and only showed a marked change during the 'dry' period of 1990 to early 1991. Density of birds in this woodland was 59.2 ± 20.8 birds/km. Biomass of birds was relatively constant throughout the study period, with marked increases in certain months when large species were present.     

Plant responses to drought and rewatering

Plants would be more vulnerable to water stress and thereafter rewatering or a cycled water environmental change, which occur more frequently under climatic change conditions in terms of the prediction scenarios. Effects of water stress on plants alone have been well-documented in many reports. However, the combined responses to drought and rewatering and its mechanism are relatively scant. As we know, plant growth, photosynthesis and stomatal aperture may be limited under water deficit, which would be regulated by physical and chemical signals. Under severe drought, while peroxidation may be provoked, the relevant antioxidant metabolism would be involved to annihilate the damage of reactive oxygen species. As rewatering, the recoveries of plant growth and photosynthesis would appear immediately through growing new plant parts, re-opening the stomata, and decreasing peroxidation; the recovery extents (reversely: pre-drought limitation) due to rewatering strongly depend on pre-drought intensity, duration and species. Understanding how plants respond to episodic drought and watering pulse and the underlying mechanism is remarkably helpful to implement vegetation management practices in climatic changing.Key words: drought stress, peroxidation, photosynthesis, relative growth rate, pre-drought limitation, rewatering, signals, stomatal conductanceUnder the climatic changing context, drought has been, and is becoming an acute problem most constraining plant growth, terrestrial ecosystem productivity, in many regions all over the world, particularly in arid and semi-arid area.^1–3 Based on the fourth assessment report by IPCC, global surface average temperature will have a 1.1–6.4°C range increase by the end of this century.³ It is indicated that a warming above 3°C would eliminate thoroughly fixed carbon function of global terrestrial vegetation, shift a net carbon source. With global warming, it is expected that water deficit would be escalated by increasing evapotranspiration, increasing the frequency and intensity of drought with an increase from 1% to 30% in extreme drought land area by 2100;³ which would offset the beneficial effect from the elevated CO₂ concentration, further limiting the structure and function of the terrestrial ecosystem. The global climate models may forecast the precipitation regimes including its distribution and amount, but the complicated responses of terrestrial ecosystem to climate change may adversely affect the predict accuracy.^1,4Plant would response to water stress by dramatically complex mechanisms from genetic molecular express, biochemical metabolism through individual plant physiological processes to ecosystem levels^2,5,6 which may mainly includes six aspects: (1) drought escape via completing plant life cycle before severe water deficit. E.g., earlier flowering in annuals species before the onset of severe drough;⁷ (2) drought avoidance via enhancing capacity of getting water. E.g., developing root systems or conserving it such as reduction of stomata and leaf area/canopy cover;^8,9 (3) drought tolerance mainly via improving osmotic adjustment ability and increasing cell wall elasticity to maintain tissue turgidity;¹⁰ (4) drought resistance via altering metabolic path for life survives under severe stress (e.g., increased antioxidant metabolism);^11,12 (5) drought abandon by removing a part of individual, e.g., shedding elder leaves under water stress;² (6) drought-prone biochemical-physiological traits for plant evolution under long-term drought condition via genetic mutation and genetic modification.^13–15 The processes may be involved in multi-aspects simultaneously in responses of plants to drought stress and thereafter rewatering.In the field context, there is always interval occurrence in drought and/or rewetting events, particular under climatic change conditions predicting more frequent drought and flooding events.³ The water cycle change may greatly impact plant growth, photosynthesis and many key metabolic functions, thereby ecosystem productivity and agricultural achievement.^5,16–18 Actually, sporadic precipitation would become a critical issue for maintaining ecosystem structural stability and even it''s surviving in arid and semi-arid area. For example, a small rainfall pulse can induce a rapid response in a desert ecosystem, which quickly triggers plant growth so that the plants can survive.¹⁹ Thus, to highlight how plant and terrestrial ecosystem cope with adverse abnormal climatic change variables is, and always will be crucial research issue in practical management of plant growth and vegetation productivity. Here, we try to provide a brief insight into how plant responses to the pre-drought and rewatering in terms of the plant growth, gas exchange and key related-physiological processes such as reactive oxygen species (ROS) metabolism. Finally a regulation path schematic is presented to try to explain the involved processes.     

Resilience of Mediterranean shrubland to a severe drought episode: the role of seed bank and seedling emergence

Extreme climate events, such as severe drought episodes, may induce changes in vegetation if they induce species‐specific adult mortality and changes in the seedling recruitment pattern. In 2005 a severe drought occurred in Doñana National Park (south Spain) causing extensive shrubland mortality. Over the following years we monitored the soil seed bank and seedling emergence via a gradient of canopy dieback induced by the drought episode. The canopy dieback corresponded to an increase in emergence of seedlings of woody species in 2007, probably because of the reduced competition induced by canopy loss. The soil seed bank of woody species sampled in 2008 was less abundant on plots with a higher proportion of dead vegetation, probably because of depletion of the seed bank as a result of the increased germination in the previous year and also as a result of a reduction in seed supply in these sites. Accordingly, in 2009 we detected reduced emergence of woody species on plots that had suffered the greatest shrub mortality. We failed to find any significant changes in patterns of the soil seed bank and seedling emergence of short‐lived herbaceous species, indicating greater resilience in these types of species. This study highlights the resilience of Mediterranean shrublands to climate fluctuations at one extreme of the variability characteristic of these ecosystems. An increase in the frequency of severe drought episodes – increasingly probable under the new climate conditions – does have the potential, however, to induce changes in vegetation, especially in woody communities that need more time to replenish their seed banks.     

Impact of drought and increasing temperatures on soil CO2 emissions in a Mediterranean shrubland (gariga)

In arid and semiarid shrubland ecosystems of the Mediterranean basin, soil moisture is a key factor controlling biogeochemical cycles and the release of CO₂ via soil respiration. This is influenced by increasing temperatures. We manipulated the microclimate in a Mediterranean shrubland to increase the soil and air night-time temperatures and to reduce water input from precipitation. The objective was to analyze the extent to which higher temperatures and a drier climate influence soil CO₂ emissions in the short term and on an annual basis. The microclimate was manipulated in field plots (about 25 m²) by covering the vegetation during the night (Warming treatment) and during rain events (Drought treatment). Soil CO₂ effluxes were monitored in the treatments and compared to a control over a 3-year period. Along with soil respiration measurements, we recorded soil temperature at 5 cm depth by a soil temperature probe. The seasonal pattern of soil CO₂ efflux was characterized by higher rates during the wet vegetative season and lower rates during the dry non-vegetative season (summer). The Warming treatment did not change SR fluxes at any sampling date. The Drought treatment decreased soil CO₂ emissions on only three of 10 occasions during 2004. The variation of soil respiration with temperature and soil water content did not differ significantly among the treatments, but was affected by the season. The annual CO₂ emissions were not significantly affected by the treatments. In the semi-arid Mediterranean shrubland, an increase of soil CO₂ efflux in response to a moderate increase of daily minimum temperature is unlikely, whereas less precipitation can strongly affect the soil processes mainly limited by water availability.     

Long‐term change within a Neotropical forest: assessing differential functional and floristic responses to disturbance and drought

Disentangling the relative roles of biotic and abiotic forces influencing forest structure, function, and local community composition continues to be an important goal in ecology. Here, utilizing two forest surveys 20‐year apart from a Central American dry tropical forest, we assess the relative role of past disturbance and local climatic change in the form of increased drought in driving forest dynamics. We observe: (i) a net decrease in the number of trees; (ii) a decrease in total forest biomass by 7.7 Mg ha^?1 but when calculated on subquadrat basis the biomass per unit area did not change indicating scale sensitivity of forest biomass measures; (iii) that the decrease in the number of stems occurred mainly in the smallest sizes, and in more moist and evergreen habitats; (iv) that there has been an increase in the proportion of trees that are deciduous, compound leaved and are canopy species, and a concomitant reduction in trees that are evergreen, simple‐leaved, and understory species. These changes are opposite to predictions based on recovery from disturbance, and have resulted in (v) a uniform multivariate shift from a more mesic to a more xeric forest. Together, our results show that over relatively short time scales, community composition and the functional dominance may be more responsive to climate change than recovery to past disturbances. Our findings point to the importance of assessing proportional changes in forest composition and not just changes in absolute numbers. Our findings are also consistent with the hypothesis that tropical tree species exhibit differential sensitivity to changes in precipitation. Predicted future decreases in rainfall may result in quick differential shifts in forest function, physiognomy, and species composition. Quantifying proportional functional composition offers a basis for a predictive framework for how the structure, and diversity of tropical forests will respond to global change.