Early post‐fire succession in northwestern Patagonia grasslands

Abstract. Large‐scale disturbances, notably fire and grazing, structure grass and shrubland dynamics in semi‐arid environments. We studied early post‐fire succession in two burned grasslands, one unburned grassland, and one shrubland near the burned area. We observed three processes: (1) establishment of a ‘phantom’ community comprised of fugitive species. Although transient, these species increase diversity and recharge the seed bank before the next disturbance; (2) regeneration of the original community by persistence of resprouter species and by auto‐replacement; (3) early stages of invasion by seedlings of the shrub Fabiana imbricata, which germinate next to shrubland and create new F. imbricata patches. Weed invasion was principally due to the ruderal exotic species Verbascum thapsus from the nearby road verge and by rapid increase of Rumex acetosella cover, another exotic species present before the fire. Although post‐fire climatic conditions are particularly important in semi‐arid environments, succession depends greatly on the regeneration strategies and dispersal abilities of the species present in the burned area. The phantom community occurs only at the first stage of succession when there is little competition for resources. We could call this process ‘the race for occupation of the area’. The second stage, when competition for resources becomes progressively more important, could be called ‘the effort to maintain space’.     

Spatial micro‐patterns in the steppe of Tierra del Fuego induced by sheep grazing

Abstract. In the Festuca gracillima tussock steppe of northern Tierra del Fuego there is a physiognomic pattern at the community scale composed of a matrix of tussocks, scrub patches of Chiliotrichum diffusum and lawn patches. In this paper we compared floristic composition, soils, microclimate and sheep use of these three types. Species composition greatly differed among physiognomic types. Lawn patches showed the highest diversity and maximum cover of exotics, grazing escape growth forms and species of high‐fertility habitats. Soil profiles of tussock steppe and lawn were similar but the A‐horizon of the latter contained more clay and cations. Radiation and temperature during daylight at the soil surface in summer were highest in the lawn patches. Soil impedance, a good indicator of trampling, was highest on the lawn patches. According to dung pellet density, lawn patches received much heavier use than the tussock matrix. Our results suggest that lawn patches supported the highest grazing impact and undergo a eutrophication process favoured by high mineral inputs, high mineralization rates in the soil during the warm season and restricted leaching through the profile. They also support the idea that a feedback mechanism between vegetation heterogeneity and sheep grazing behaviour maintains the patches. Although grazing behaviour suggests that the lawn patches favour animal production, their expansion would result in a system less protected against the impact of water deficits and temperature variations and more vulnerable to erosion and to the loss of important species.     

Interactive effects of introduced herbivores and post‐flowering die‐off of bamboos in Patagonian Nothofagus forests

Question: In November 2000, Chusquea culeou, a bamboo species dominating Andean forest understories in southern Argentina and Chile, massively flowered and died over a north‐south distance of ca. 120 km. Because bamboo is the major forage for large herbivores in these forests, we examined the interactive influences of the bamboo die‐off and herbivory by introduced cattle on understory and tree regeneration. Location: Lanín National Park, Argentina. Methods: Permanent plots, in and outside livestock exclosures, were installed in a Nothofagus dombeyi forest in patches of flowered and non‐flowered C. culeou. Plots were monitored over four years for changes in understory composition and tree seedling densities and heights. Results: After the C. culeou die‐off, new establishment of N. dombeyi was low, both with and without herbivory. Livestock alone directly increased N. dombeyi seedling mortality through physical damage. However, tree seedling browse ratings and height growth were interactively affected by bamboo flowering and herbivory; unfenced plots in flowered bamboo patches had the shortest seedlings, highest browse ratings, and lowest tree seedling annual growth rates. Understory cover was higher where livestock were excluded, and this effect was intensified in the patches of flowered bamboo. Neither herbivory nor bamboo flowering resulted in major changes in species composition, with the exception of Alstroemeria aurea. Conclusion: Effects of livestock on N. dombeyi regeneration were contingent on flowering of C. culeou. Prior to introduction of livestock, N. dombeyi regeneration was probably successful beneath canopy gaps during windows of opportunity following bamboo die‐off, but now livestock impede tree regeneration. Herbivory during bamboo withering periods also produces more open understories, particularly affecting palatable heliophyllous herb species such as Alstroemeria aurea. The results underscore the importance of assessing herbivore impacts on tree regeneration during relatively short periods of potential tree regeneration immediately following rare bamboo flowering and die‐off.     

Influence of nests of leaf‐cutting ants on plant species diversity in road verges of northern Patagonia

Abstract. It has been suggested that ant nests are the most frequent small‐scale disturbance that affect vegetation patterns. However, their effects on plant diversity are little studied. We document effects of nests of the leaf‐cutting ant Acromyrmex lobicornis on physical‐chemical soil properties and their influence on plant diversity near road verges in a desert steppe in NW Patagonia, Argentina. We analysed nest soils and controls for nitrogen, phosphorus, organic matter, moisture retention capacity and texture. We also analysed the vegetation on 42 nests (30 active and 12 abandoned or without life) and 42 areas without nests. Soil around nests had a greater nutrient content and capacity to retain moisture than control soils, which is mainly due to the presence of organic waste that the ants deposit on the soil surface. We found no association between the occurrence of nests and specific groups of plants, but plant diversity was higher at nest‐sites than at nearby non‐nest sites. This increased diversity – which is also found on abandoned nests – is mainly due to the occurrence of a larger number of native and exotic plant species on nest‐sites that are uncommon elsewhere in the study area. The most abundant plant species showed similar cover values at nest and non‐nest sites. This suggests that changes in diversity are associated to edaphic changes caused by nests rather than by changes in competitive balance caused by dominant species exclusion. We propose that the nests of Acromyrmex lobicornis, through increasing the availability of resources, generate favourable microsites that can function both as ‘refuges’ for less frequent native species, and as‘stepping stones’ for less frequent exotic plant species.     

Sex‐related spatial patterns of Poa ligularis in relation to shrub patch occurrence in northern Patagonia

Abstract. Poa ligularis is a dioecious species and a valuable forage plant which is widespread in the arid steppe of northern Patagonia (Argentina). The vegetation in these areas consists of a system of perennial plant patches alternating with bare soil areas defining contrasting micro‐environments. We hypothesized that (1) male and female individuals of P. ligularis are spatially segregated in different micro‐environments, (2) the intensity of spatial segregation of sexes depends on plant structure and (3) spatial segregation of sexes is enhanced by competitive interactions between the sexes within the vegetation patches. We analysed the spatial distribution of female and male individuals in relation to the spatial pattern of vegetation in two areas differing in their vegetation structure. The location of P. ligularis within patches where either male, female or both sexes occurred was also analysed. The results indicate that different patterns of spatial distribution of sexes of P. ligularis may be found at the community level depending on the dominant life forms and geometric structure of plant patches. Where patches are of a lower height, with a high internal patch cover, individuals of both sexes are concentrated within patch canopies. In sites characterized by large, tall patches and less internal patch cover suitable microsites for female and male P. ligularis occur both within and outside the patch with males located at further distances from the patch edge. Where the patch is large and tall enough to allow the establishment of males and females at relatively high numbers, males occupy the patch periphery or even colonize the interpatch bare soil. These spatial patterns are consistent with selective traits in which females better tolerate intraspecific competition than males, while males tolerate wider fluctuations in the physical environment (soil moisture, nitrogen availability, wind intensity, etc.).     

Plant functional types and ecosystem function in relation to global change

Abstract. Plant functional types (PFTs) bridge the gap between plant physiology and community and ecosystem processes, thus providing a powerful tool in climate change research. We aimed at identifying PFTs within the flora of central-western Argentina, and to explore their possible consequences for ecosystem function. We analyzed 24 vegetative and regenerative traits of the 100 most abundant species along a steep climatic gradient. Based on plant traits and standard multivariate techniques, we identified eight PFTs. Our results confirmed, over a wide range of climatic conditions, the occurrence of broad recurrent patterns of association among plant traits reported for other floras; namely trade-offs between high investment in photosynthesis and growth on the one hand, and preferential allocation to storage and defence on the other. Regenerative traits were only partially coupled with vegetative traits. Using easily-measured plant traits and individual species cover in 63 sites, we predicted main community-ecosystem processes along the regional gradient. We hypothesized likely impacts of global climatic change on PFTs and ecosystems in situ, and analysed their probabilities of migrating in response to changing climatic conditions. Finally, we discuss the advantages and limitations of this kind of approach in predicting changes in plant distribution and in ecosystem processes over the next century.     

Effects of topography,soil moisture,wind and grazing on Festuca seedlings in a Patagonian grassland

Abstract. The effects of topography, soil moisture, wind and grazing on the emergence and survival of seedlings of Festuca spp. were examined in the steppe zone of Patagonia, Argentina. Ungrazed and grazed field treatment plots were established on a plain and a north-facing slope at the Media Luna Ranch (43° 36′S, 71° 25′W). On the leeward and windward sides of each of 15 Festuca plants, 0.1 m × 0.4 m quadrats were censused bimonthly for seedling emergence and survival over three growing seasons. Three categories were distinguished: recently germinated and up to the first leaf, two to four leaves, and from five leaves up to one tiller. Soil moisture content, litter cover and frost heaving effects were also determined for each treatment at each sampling date. Festuca spp. showed two emergence peaks, one in late fall and the other in early to mid-spring. Seedling emergence was significantly correlated with soil moisture content in the 0–5 cm of the soil during the three growing seasons. Seedlings that emerged in the fall had higher survivorship than those that emerged in spring. Seedling emergence and survival was significantly (p < 0.01) lower on slopes, in the grazing treatment, and on windward sides of adult plants. In this grassland, an increase in the availability of safe sites for seedling emergence and survival might be achieved by protecting vegetation from grazing, particularly on north-facing slopes.     

Seedling recruitment in a semi‐arid Patagonian steppe: Facilitative effects of refuse dumps of leaf‐cutting ants

Question: What is the influence of refuse dumps of leaf‐cutting ants on seedling recruitment under contrasting moisture conditions in a semi‐arid steppe? Location: Northwestern Patagonia, Argentina. Methods: In a greenhouse experiment, we monitored seedling recruitment in soil samples from refuse dumps of nests of the leaf‐cutting ant Acromyrmex lobicornis and non‐nest sites, under contrasting moisture conditions simulating wet and dry growing seasons. Results: The mean number of seedling species and individuals were higher in wet than in dry plots, and higher in refuse dump plots than in non‐nest soil plots. The positive effect of refuse dumps on seedling recruitment was greater under low moisture conditions. Both the accumulation of discarded seeds by leaf‐cutting ants and the passive trapping of blowing‐seeds seems not explain the increased number of seeds in refuse dumps. Conversely, refuse dumps have higher water retention capacity and nutrient content than adjacent non‐nest soils, allowing the recruitment of a greater number of species and individual seedlings. Conclusions: Nests of A. lobicornis may play an important role in plant recruitment in the study area, allowing a greater number of seedlings and species to be present, hence resulting in a more diverse community. Moreover, leaf‐cutting ant nests may function as nurse elements, generating safe sites that enhance the performance of neighbouring seedlings mainly during the driest, stressful periods.     

Interactive effects of fire and grazing on structure and diversity of Mediterranean grasslands

Abstract. The separate and combined effects of fire and cattle grazing on structure and diversity of productive Mediterranean grasslands in northern Israel were examined within a set of climatically and edaphically similar sites. Cover and height of green and dry plants in winter, and species richness and diversity in spring, were measured in paired transects on both sides of cattle fences, and on both sides of boundaries of both incidental and experimentally lit fires. Early in the first growing season after a fire, plant cover as well as height of green plants were reduced, compared to unburnt grassland. These structural effects of fire were similar to the effects of grazing, but they were greater in ungrazed than in grazed grasslands, indicating a fire-grazing interaction. The effects of fire were considerably attenuated in the second growing season after the fire. Species richness and diversity tended to be higher in grazed than in adjacent ungrazed grasslands. Richness consistently increased after a fire only in grazed grasslands with a strong perennial component. In ungrazed grasslands, and in predominantly annual grasslands, fires reduced species richness and diversity at least as often as they increased it. Fire and grazing should be regarded as two agents with distinct and interactive effects on the community, rather than as two alternative mechanisms of a general disturbance factor.     

Facilitation by nurse shrubs of resprouting behavior in a post-fire shrubland in northern Patagonia,Argentina

Abstract. The nurse-plant syndrome is a widely recognized example of positive (facilitative) influences of plant species on the establishment and growth of other species. Most studies of the nurse-plant syndrome have been on species that reproduce mainly from seed rather than vegetatively. In this study, we experimentally compared the influences of two species of nurse shrubs, Schinus patagonicus and Berberis buxifolia, on the survival and growth of vegetatively reproducing herbaceous and woody plants in a post-fire shrubland in northern Patagonia, Argentina. The vegetation beneath shrubs was removed by clipping and, in a paired-sample design, one half of the canopy of each shrub was removed. We determined species richness, counted number of resprouts, and measured photon flux density and soil moisture beneath cut and uncut halves of each shrubs. Abundances of resprouts were several times greater beneath the uncut vs. the cut shrubs, as was the mean number of species. Thus, shrubs have a strong facilitating influence as measured by resprout densities and the number of species. Numbers of resprouts and of species were twice as high beneath Schinus as beneath Berberis, implying important differences in the facilitative effects of the two shrubs species. Microsites beneath Schinus were characterized by lower and more heterogeneous light levels but by greater soil moisture. Even though the reproductive mode in this post-fire shrubland is overwhelmingly vegetative, facilitation by nurse shrubs is important and differentially effective for different species of nurse shrubs.     

Plant functional traits and environmental filters at a regional scale

Abstract. Links between plant traits and the environment, i.e. sets of plant attributes consistently associated with certain environmental conditions, are the consequence of the filtering effect of climatic, disturbance and biotic conditions. These filters determine which components of a species pool are assembled into local communities. We aimed at testing for consistent association between plant traits and climatic conditions along a steep regional gradient, divided into 13 climatically homogeneous sectors, in central-western Argentina. We analyzed 19 vegetative and regeneration traits of the 100 most abundant species along the gradient. For each trait, we tested for homogeneity of frequencies of different categories between sectors and the regional species pool, using the χ² statistic. We rejected H₀ in 71% of the pair-wise comparisons, which strongly suggests a ‘filtering effect’ of climatic factors on key plant functions. Vegetative traits were filtered more often than regeneration traits. Specific leaf area, life span, ramification, canopy height, leaf weight ratio, carbon investment into support tissue and pollination mode were the traits showing differences in the largest number of pair-wise comparisons. This is probably the first attempt to detect, on a quantitative, statistically conservative basis, consistent linkages between climatic factors and numerous plant traits, over a broad spectrum of environmental conditions and plant growth forms. We discuss the advantages and limitations of this approach in predicting vegetation structure and functioning under present environmental conditions, and those expected for the next century as a consequence of global change.     

Vegetation heterogeneity and diversity in flat and mountain landscapes of Patagonia (Argentina)

Abstract. We studied floristic and diversity patterns and their environmental controls in two landscapes of contrasting topography in the Patagonian steppe. The analyses were focused on the effects of water availability gradients and landscape configuration on plant species distribution and coexistence. Floristic variation was investigated using Correspondence Analysis. The relationship between floristic and environmental variation was analyzed using Canonical Correspondence Analysis and correlation tests. We explored diversity patterns by relating spatial distance to floristic dissimilarities. The floristic gradient was determined by shrub and grass species and was related to precipitation in the flat area, and to precipitation, elevation and potential radiation in the mountain area. Site species richness increased with water availability in both areas. Mean site species richness and species turnover in space was higher in the mountain than in the flat area. Landscape species richness and floristic gradients were more concentrated in the mountain than in the flat area. In contrast to shrubs and grasses, forb species distributions were uncoordinated and probably independent of any environmental gradient. Our results suggest (1) that landscape configuration affects species composition and diversity through its direct effect on abiotic environmental heterogeneity, and (2) that the environmental controls of the community composition vary depending on the plant functional type considered.     

Vertical stratification of ichneumonid wasp communities: the effects of forest structure and life‐history traits

Parasitoid wasp communities of the canopy of temperate forests are still largely unexplored. Very little is known about the community composition of parasitoids between canopy and understory and how much of this difference is related to forest structure or parasitoid biological strategies. In this study we investigated upon the difference in the community composition of the parasitic wasps Ichneumonidae between canopy and understory in a lowland temperate forest in northern Italy. We used general linear models to test whether parasitic strategy modifies species vertical stratification and the effect of forest structure. We also tested differences in β‐diversity between canopy and understory traps and over time within single forest layers. We found that stand basal area was positively related to species richness, suggesting that the presence of mature trees can influence local wasp diversity, providing a higher number of microhabitats and hosts. The ichneumonid community of the canopy was different from that of the understory, and the β‐diversity analysis showed higher values for the canopy, due to a higher degree of species turnover between traps. In our analyses, the vertical stratification was different between groups of ichneumonids sharing different parasitic strategies. Idiobiont parasitoids of weakly or deeply concealed hosts were more diverse in the understory than in the canopy while parasitoids of spiders were equally distributed between the two layers. Even though the ichneumonid community was not particularly species‐rich in the canopy of the temperate forests, the extension of sampling to that habitat significantly increased the number of species recorded.     

Floristic composition of a Swedish semi‐natural grassland during six years of elevated atmospheric CO2

Abstract. A semi‐natural grassland in Sweden was exposed to an elevated CO₂ concentration during a six‐year open‐top chamber experiment. Vegetation composition was assessed twice a year using the point‐intercept method. The field had been grazed previously, but when the experiment started this was replaced with a cutting regime with one cut (down to ground level) each year in early August. From the third to the sixth year of the study the harvested material was divided into legumes, non‐leguminous forbs and grasses, dried and weighed. Elevated CO₂ had an effect on species composition (as analysed by Principal Component Analysis) that increased over time. It also tended to increase diversity (Shannon index) in summer, but reduce it in spring. However, the effects of the weather and/or time on species composition and diversity were much more prominent than CO₂ effects. Since the weather was largely directional over time (from dry to wet), with the exception of the fifth year, it was difficult to distinguish between weather effects and changes caused by a changed management regime. In all treatments, grasses increased over time in both mass and point‐intercept measurements, whereas non‐leguminous forbs decreased in mass, but not in point‐intercept measurements. Legumes increased in the point‐intercept measurements, but not in biomass, at elevated CO₂, but not in the other treatments. Overall, we found that elevated CO₂ affected species composition; however, it was only one of many factors and a rather weak one.     

Biodiversity of semi‐arid Mediterranean grasslands: Impact of grazing and afforestation

Question: What is the impact of grazing and/or afforestation on grassland diversity, species composition and cover parameters? Location: Semi‐arid Mediterranean grasslands of Jordan. Methods: Vegetation, litter, bare soil and rock cover were compared among four management types – free grazing and protected from grazing with three levels of tree cover. Species composition, plant cover, species richness and evenness were used to evaluate differences in vegetation among management types. Species composition differences among management types were also investigated. Results: Semi‐arid Mediterranean grasslands harbour appreciable levels of plant biodiversity. Grazing did not affect plant diversity, indicating the high resilience against and adaptation to grazing; however,grazing affected species composition and cover parameters. Afforestation seems to protect soil through higher litter cover but its impact on plant biodiversity was negative and markedly affected species composition. Conclusions: Neither protection from grazing or massive afforestation alone are sufficient for conserving biodiversity in this system. A management model is suggested where the landscape should be maintained as a mosaic of four management types: complete protection from grazing, grazing rotation, planting sparse trees in eroded areas and revegetating degraded areas using native, herbaceous and grazing tolerant species.     

Species associations and nurse plant effects in patches of high-Andean vegetation

Abstract. In high-elevation communities of the southern Andes, plant cover is low due to severe environmental conditions and vegetation occurs mostly as isolated small (< 1 m²⁾ patches. Most patches are dominated by flat cushion plants. We evaluated patterns of plant species co-occurrence and species affinity for patches with and without cushion plants and different species richness. We mapped and recorded species composition of patches occurring within two 20 m × 20 m plots at the NE slope of Cerro Chall-Huaco, Nahuel Huapi National Park, Argentina. In these plots, we identified 32 and 24 plant species, and a maximum of 15 and 12 species per patch, respectively. The community was characterized by positive associations between species. Patches in which either of the common cushion plants Mulinum leptacanthum and Oreopolus glacialis occurred sustained richer communities than patches in which they were absent. Patches dominated by different cushion plants did not differ in species composition, but species differed in their affinities for patches with different numbers of species. Because richness increased with patch size and patch size with time, differential affinities of plant species suggest that successional changes take place in the patches. Some small herbaceous species appear to act as late colonizers, mostly restricted to species-rich patches. Flat cushion plants are considered ‘nurse plants’; they strongly modify micro-environmental conditions and allow establishment and survival of associated species.     

Vegetation‐environment relationships in Atlantic European calcareous grasslands

Abstract. The relationship between vegetation and environment was investigated for calcareous grasslands in a region in the west of Spain, France, Britain and Ireland defined by climatic criteria. Vegetation was sampled using objective methods and data collected on soils, land cover, location and management. Climate data were obtained from an available database. Examination of the first axis of vegetation variation as defined by Detrended Correspondence Analysis (DCA) showed a gradient from the Irish and British samples to those from France. The Spanish samples formed a separate group on the second axis. The species composition along the gradients is discussed. Correlations between the vegetation gradients and environmental variables were determined. The strongest correlations with the first DCA axis were for temperature, latitude, soil organic matter, grazing and land cover. The second DCA axis was highly correlated with rainfall, altitude and land cover. The third and fourth DCA axes were more difficult to interpret but appeared to be related to land cover. The results indicate that climate factors are important at this scale, but should not be considered in isolation and that factors relating to land cover and management should also be taken into account.     

Site quality effects on post‐fire regeneration of Pinus brutia forest on a Greek island

Abstract. Natural stands of Pinus brutia were compared to burned areas after a great fire in 1985 on the island of Thasos, North Greece, in relation to site quality effects on the success of natural regeneration and understorey vegetation. The number of seedlings in the natural stands of P. brutia was 1124/ha against 3188/ha in the burned area. In both cases natural regeneration was successful. Under the canopy of P. brutia forests the predominant shrubs were Quercus coccifera and vPhillyrea latifolia. A high percentage (up to 40%) of the total area was covered by these species. In the burned areas, 12 yr after the wildfire of 1985, Cistus incanus and C. salvifolius played a significant role in the survival of P. brutia seedlings. These species had covered the burned area long before the pine seed germination.     

Restriction of transpiration rate under high vapour pressure deficit and non‐limiting water conditions is important for terminal drought tolerance in cowpea

Drought stress is a major constraint on cowpea productivity, since the crop is grown under warm conditions on sandy soils having low water‐holding capacity. For enhanced performance of crops facing terminal drought stress, like cowpea, water‐saving strategies are crucial. In this work, the growth and transpiration rate (TR) of 40 cowpea genotypes with contrasting response to terminal drought were measured under well‐watered conditions across different vapour pressure deficits (VPD) to investigate whether tolerant and sensitive genotypes differ in their control of leaf water loss. A method is presented to indirectly assess TR through canopy temperature (CT) and the index of canopy conductance (Ig). Overall, plants developed larger leaf area under low than under high VPD, and there was a consistent trend of lower plant biomass in tolerant genotypes. Substantial differences were recorded among genotypes in TR response to VPD, with tolerant genotypes having significantly lower TR than sensitive ones, especially at times with the highest VPD. Genotypes differed in TR response to increasing VPD, with some tolerant genotypes exhibiting a clear VPD breakpoint at about 2.25 kPa, above which there was very little increase in TR. In contrast, sensitive genotypes presented a linear increase in TR as VPD increased, and the same pattern was found in some tolerant lines, but with a smaller slope. CT, estimated with thermal imagery, correlated well with TR and Ig and could therefore be used as proxy for TR. These results indicate that control of water loss discriminated between tolerant and sensitive genotypes and may, therefore, be a reliable indicator of terminal drought stress tolerance. The water‐saving characteristics of some genotypes are hypothesised to leave more soil water for pod filling, which is crucial for terminal drought adaptation.