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.     

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.).     

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’.     

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.     

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.     

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.     

Vertical structure of wet grasslands under grazed and non‐grazed conditions in Tierra del Fuego

Abstract. We studied the vertical structure of wet grazed grasslands in Tierra del Fuego (southern Argentina). A point quadrat method was developed using a fine needle graduated in cm. The vertical and horizontal frequency of species and organs was quantified in samples collected from non‐grazed and grazed plots in the field. There was vertical stratification in both types of samples, but only in the first eight cm above the ground in grazed samples, with a dominance of Caltha sagittata. In non‐grazed samples graminoids grew taller than forbs and their inflorescences were an important element of the canopy structure. In both treatments, vertical species diversity was maximum in the lower part of the canopy, although diversity was significantly higher in grazed pots. In grazed samples, Caltha sagittata was the dominant species in 46% of samples and its leaves occupied 35% of the upper canopy. In the non‐grazed samples, Hordeum pubiflorum and Festuca magellanica were dominant in 63% of the samples, with H. pubiflorum leaves occupying 55% of the upper canopy. Comparing species by pairs, significant differences in vertical position were maintained in non‐grazed versus grazed pots. It is concluded that vertical stratification occurs even in the shortest communities. In this community, forb species grew close to the ground in the grazed areas, while forbs grew in the gaps and grasses above them in the non‐grazed areas. The main differences were in the relative dominance of forb and grass species and the presence of inflorescences.     

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.     

Early post-fire regeneration of a Pinus halepensis forest on Mount Párnis,Greece

Abstract. The post-fire regeneration of a 45-yr-old Pinus halepensis (Aleppo pine) forest, burned in July 1989, has been studied on Mount Párnis, Attiki, Greece. Four experimental plots at various slopes and exposures were established at altitudes of 400 - 450 m, and monitored for 3 yr at 3-month intervals. Early regeneration took place abundantly, through both resprouting and seed germination of mostly hard-seeded herbs and shrubs; the floristic richness was high with 80 taxa. Pine seedling emergence took place during the winter of the first post-fire year. The mean pine seedling density by the end of the recruitment period (March 1990) was 5–6 seedlings/m². This density decreased slightly during late spring and considerably during summer. During the second post-fire year only a relatively slight decline was observed; thereafter the density was stabilized to 1 - 2 seedlings/m². Mortality follows a negative exponential curve that levels off at ca. 20 %. Height distributions throughout the three post-fire years were all positively skewed as a result of the presence of few very tall saplings. A considerable fraction (20 %) of very short (5–15 cm) saplings were still alive 39 months after the fire; these may constitute the sapling bank. Based on the analysis of height distribution curves, it is concluded that the taller seedlings survived significantly better than the shorter ones.     

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.     

Seasonal variation in the seed bank of a Patagonian grassland in relation to grazing and topography

Abstract. The germinable seed bank of Festuca spp., Carex patagonica, and other annuals and perennials in a semiarid Patagonian grassland was analyzed every three months. The effects of grazing, topography and microsites with respect to established grass tussocks on the germinable-seed bank were also analyzed. The total germinable-seed bank was larger in summer after seed rain. At this time of the year most of the seeds were from annuals and perennial dicots. Seeds of the dominant Festuca spp. were the main components of the graminoid seed bank, which was homogeneously distributed in patches of bare soil. In spring, i.e. some months after the seed rain, the germinable-seed bank of most of the species was greatly reduced, while the seed bank of Carex patagonica did not change significantly. At this time of the year, the graminoid seed bank was heterogeneously distributed in space, with the seeds accumulating in wind-protected microsites. In the grazing treatments, the seed bank of the perennial grasses was reduced proportionally to the decrease of the plant cover. On the other hand, the germinable-seed bank of C. patagonica increased with the grazing treatments, in relation to the increase of vegetation cover. Topographical position had two types of effect on the seed bank of perennial grasses. One was the increase of the germinable-seed bank, after the seed rain, on the relatively warmer northeast facing slopes, which was related to a higher seed rain in these places. The other was an increased seed loss on slopes. The different seed-bank strategies observed in this grassland are compared and discussed in relation to strategies described for other grasslands.     

Spatial distribution of Tephrocactus aoracanthus (Lem.) Lem. in relation to shrubs in the hyperarid regions of west‐central Argentina

The spatial intraspecific and interspecific distribution of Tephrocactus aoracanthus (Lem.) Lem. was studied using univariate and bivariate point pattern analysis, at landscape scale in four different physiographic units and at microscale in terms of the shrubs found in those units. The influence of shrubs on photosynthetically active radiation and on soil temperature under their canopies was analysed. Results show a high environmental variability in light and temperature under shrubs that generally has no influence on the distribution of T. aoracanthus. Apparently, this cactus has great ability to adapt to different conditions of both photosynthetically active radiation and temperature. However, intraspecific distribution was highly clustered in all physiographic units, independently of bushes. Possibly, the mode of seed dispersal and the rooting of cladodes at small scale are the main processes that determine the strong intraspecific clustering of T. aoracanthus.     

Invasibility of a coastal strip in NE Spain by alien plants

Abstract. The alien plant species and the ecological factors that facilitate their invasion to a coastal strip in the Baix Camp region (Tarragona, NE Spain) were studied. A detailed inventory of the area showed that 20% of the plant species, most of them from the American Continent, were aliens, many of which were strongly invasive. At the habitat level, the relationships between the invasive behaviour and a number of autecological, ecological and habitat variables were analysed by means of logistic analyses. In the best model five variables were related to alien plant invasions: climatic affinity, disturbance of the invaded habitat, route of introduction, soil moisture during summer, and life form. To elucidate the environmental factors that could be responsible for alien plant invasions at a regional scale, the number of alien taxa and the environmental features at 13 localities of the western Mediterranean area were analysed. The warm lowlands, subjected to low annual rainfall and high population density, exhibited the greatest reception capacity for alien plants. It is concluded that various specific environmental features – high mean temperatures, the abundance of riparian systems, the existence of biotopes with favourable water balance which harbour a great richness of alien species and the high human pressure to which the enclave is subjected, combined with the ecological requirements of the alien species and certain species attributes (biotype) can interact at different scales and have caused the over‐representation of alien plants.     

Dynamics of understory vegetation in an old-growth boreal coniferous forest, 1988–1993

Abstract. Understorey vegetation changes in a South Norwegian old-growth coniferous forest were studied between 1988 and 1993 in 200 1-m² vegetation plots. Our aims were to quantify the amount of between-year compositional change, and to elaborate the environmental basis for long-term vegetation change, including the previously identified gradient structure with a major gradient related to topography (and soil nutrient status and soil depth) and a minor gradient reflecting paludification and canopy coverage. Species richness (yearly mean and cumulative species number) and change in species richness differed between vascular plants and cryptogams, and between forest types. The number of vascular plant species decreased in pine forest in dry years; bryophyte species number increased in spruce forest. Statistically significant vegetation change, as tested by constrained ordination (CCA) with time as the constraining variable, is demonstrated for most one-year periods and for the five-year period in most forest types. Vegetation change along identified gradients, measured as plot displacement along DCA ordination axes, also occurred. The magnitude of year-to-year vegetation change was related neither to forest type nor to one-year period; different responses to climatic and environmental change were observed in each forest type. The largest average displacement observed, from medium-rich spruce forest towards poor spruce forest, was interpreted as a long-term trend. Humus-layer pH decreased by ca. 0.25 units from 1988 to 1993, most strongly in medium-rich spruce forest where exchangeable Ca decreased and Al and Mn increased strongly. Our study supports the hypothesis that vascular plants show a long-term and broad-scale response to soil acidification. Change in bryophyte composition is linked to some very long growing-seasons. Detailed analysis of short-term vegetation dynamics enhances the interpretation of long-term changes and stresses the complementarity of univariate and multivariate methods in the analysis of vegetation change.     

Effects of selective logging on tree diversity,composition and plant functional type patterns in a Bornean rain forest

Abstract. The effects of selective logging on tree diversity, changes in tree species composition and plant functional types were studied with the use of seven permanent plots in virgin and in logged forest. All plots were located in a lowland dipterocarp rain forest in East Kalimantan on the island of Borneo. Just after logging and during the following 20 yr tree diversity measured as Fishers’α was not significantly affected in logged forest plots. Temporal shifts in tree species composition were analysed with Principal Component Analysis (PCA). Logged forest plots had much larger changes over time than virgin forest plots. In the smallest diameter class, some logged forest plots showed a distinct trajectory in PCA space compared to virgin forest plots, while in larger diameter classes movement of logged plots in PCA space was random. This suggests that there is no predetermined community to which logged forest plots tend to shift when recovering from logging. We found a significant negative correlation between diameter increments and the species‐specific wood densities of tree species. Species‐specific wood density and potential tree height were used to assign species to five PFTs. As expected, logging increased the fraction of softwood stems in small diameter classes. In the largest diameter classes (≥ 50 cm DBH) a strong decrease of softwood emergent stems was found in logged forest plots. After more than 20 yr no recruitment was found of softwood emergent stems in selectively logged forest.     

Relative importance of abiotic and land use factors in explaining variation in woody vegetation in a French rural landscape

Abstract. In a rural landscape, scale vegetation patterns of woody species are controlled by both abiotic and land use factors. The woody species composition in 126 sample plots was analysed and land use factors and some abiotic parameters were quantified using land register data. The relative importance of land use and abiotic factors was differentiated using a partial Canonical Correspondence Analysis (CCA); the influence of land use in neighbouring areas was explored by changing the scale of land use sampling. The woody species composition appears to be controlled equally by land use and abiotic factors. The fraction of floristic variation in tree species composition explained by land use variables was 33.2 % in 1980 and 30.8 % in 1992, while abiotic variables accounted for 31.2 %. Part of the 17 % of the floristic variance explained is related to the surrounding land use. Thus, when the land use of the sampled plots and the surrounding land use are considered simultaneously, up to 36.9 % of the species variation may be explained. Partial CCA enabled us to quantify the respective proportion of floristic variance which could be explained by land use (36.9 %), abiotic variables alone (20.2 %), shared variance (12.0 %) and unexplained variance (31.2 %). Our results indicated that a delayed effect of variation in land use on plant populations may exist. This delay may result either from population characteristics or from inadequate land use assessment. This study indicates the need for simultaneously examining land use and abiotic patterns in ecological studies, as many Mediterranean-type ecosystems have been shaped by these patterns.     

High‐throughput method for the analysis of venlafaxine in pharmaceutical formulations and biological fluids,using a tris(2,2′‐bipyridyl) ruthenium(II)–peroxydisulphate chemiluminescence system in a two‐chip device

A simple, rapid and sensitive chemiluminescent (CL) method for the assay of venlafaxine (VEN) in pharmaceutical formulations and serum samples by a two‐chip device is proposed. The method is based on the reaction of this drug with a tris(2,2′‐bipyridyl) ruthenium(II)–peroxydisulphate CL system. The optimum chemical conditions for CL emission were investigated. The calibration graph was linear for the concentration range 0.02–8.0 µg/mL. The detection and quantification limits were found to be 0.006 and 0.018 µg/mL, respectively, while the relative standard deviation (RSD) was <2.0%. The present CL procedure was applied to the determination of VEN in pharmaceutical formulations and serum samples; the recovery levels were in the range 96.5–101.2%. The results suggest that the method is unaffected by the presence of common formulation excipients found in these samples. Copyright © 2012 John Wiley & Sons, Ltd.     

The effect of elevated hydrostatic pressure upon selected biomarkers in juvenile blackspot seabream Pagellus bogaraveo in a 14 day‐long experiment

Hydrostatic pressure elevated to 500 kPa for 14 days was found to affect hepatic 7‐ethoxyresorufin‐O‐deethylase (EROD), oxidized protein (POx), protein yield and branchial Na⁺–K⁺‐ATPase. No effect on glutathione‐S‐transferase (GST), superoxidase dismutase (SOD), catalase (CAT), lipid peroxidation (LP), acetylcholinesterase (AChE), butyrylcholinesterase (BChE), condition factor (K) and hepato‐somatic index (I_H) was encountered.