Leaf growth dynamics in four plant species of the Patagonian Monte, Argentina

Studying plant responses to environmental variables is an elemental key to understand the functioning of arid ecosystems. We selected four dominant species of the two main life forms. The species selected were two evergreen shrubs: Larrea divaricata and Chuquiraga avellanedae and two perennial grasses: Nassella tenuis and Pappostipa speciosa. We registered leaf/shoot growth, leaf production and environmental variables (precipitation, air temperature, and volumetric soil water content at two depths) during summer-autumn and winter-spring periods. Multiple regressions were used to test the predictive power of the environmental variables. During the summer-autumn period, the strongest predictors of leaf/shoot growth and leaf production were the soil water content of the upper layer and air temperature while during the winter-spring period, the strongest predictor was air temperature. In conclusion, we found that the leaf/shoot growth and leaf production were associated with current environmental conditions, specially to soil water content and air temperature.     

Effect of grazing on plant patterns in arid ecosystems of Patagonian Monte

Our objective was to assess the relationship between the spatial patterning of perennial grasses (total, grazed, and non‐grazed) and shrub patches in rangelands under different grazing pressures of the Patagonian Monte. We selected three grazed paddocks with the usual stocking rate for the area, where previous studies showed that a piosphere formation is common. At each paddock, we analysed the grain of heterogeneity at sites located at two distances from the single watering point (near, far), using high‐resolution aerial photographs. At these sites, we also assessed in the field the density, size, cover, and spatial patterning of grazed and non‐grazed perennial grasses and shrub patches. The grain of heterogeneity of shrub patches was coarser in sites near the watering point than in those distant from it, as a consequence of the increase in size of both, bare soil and shrub patches. Field sampling showed that a coarse grain of heterogeneity relative to fine‐grained sites resulted from changes in species composition, increased bare soil areas and reduced perennial grass cover. In coarse‐grained sites, lower perennial grass cover resulted from lower density and/or smaller size of grass bunches than in fine‐grained sites. We did not find significant differences among sites in the proportion of perennial grazed grasses. Since the density and cover of perennial grasses was higher in fine‐ than in coarse‐grained sites, we suggested that fine‐grained sites are more important as feeding stations than coarse‐grained sites. The consequences of this differential use could lead to degradation of fine‐grained sites and to higher homogeneity in spatial plant structure and floristic composition within paddocks with respect to the condition observed at present, increasing the size of the highly degraded zone within the piosphere. At the patch level, we found that at about one third of the sampled transects, both total and non‐grazed perennial grasses were spatially aggregated with shrub patches. However, in most transects grazed perennial grasses were indifferently distributed in relation with shrub patches, showing that grazers display high selectivity of foraging sites at macro level (i.e. high and low grazing pressure sites at the paddock level), but random occupancy of vegetation units (randomness in the distribution of grazed perennial grasses at the patch level). The intensity of the positive association between non‐grazed grasses and shrub patches was higher in fine‐grained than in coarse‐grained sites and may be attributed to higher protection against herbivores associated to denser shrub patches in fine‐ relative to coarse‐grained sites. We concluded that a feedback exists between the spatial distribution of species preferred by grazers and the spatial patterning of use of these species.     

Leaf litterfall, fine-root production, and decomposition in shrublands with different canopy structure induced by grazing in the Patagonian Monte, Argentina

Selective sheep grazing in the Patagonian Monte induces the reduction of total and perennial grass cover, species replacement within life forms, and the increase in dominance of long-lived evergreen woody plants with slow growth rates and high concentration of secondary compounds in leaves. We hypothesized that these changes in the canopy structure induced by sheep grazing will affect the mass, chemistry and decomposability of leaf litter and fine roots. We selected two sites in the Patagonian Monte, representative of ungrazed and grazed vegetation states. At each site, we assessed canopy structure (total cover and absolute and relative grass and shrub cover), monthly leaf litterfall, and fine-root biomass and production in the upper soil (15 cm). We also estimated the rates of mass, C, soluble phenolics, lignin and N decay in litterbags containing both leaf litter and fine roots of each site under field conditions during two consecutive years. The ungrazed site exhibited higher total plant cover, absolute and relative grass- and shrub-cover than the grazed one. Leaf litterfall was lower at the grazed site than at the ungrazed site. Fine-root production did not vary between sites. Leaf litter and fine root tissues had higher concentration of secondary compounds at the grazed than at the ungrazed site. However, fine roots showed lower mass and C decay than leaf litter, attributable to the predominant secondary compound (lignin and soluble phenolics, respectively). Leaf litter decomposed slower but released more N during decay at the ungrazed than at the grazed site, probably due to its low concentration of secondary compounds. We concluded that changes in canopy structure induced by grazing disturbance such as those explored in our study could reduce leaf litterfall mass and increase the concentration of secondary compounds of both leaf litter and fine roots leading to slow N release to soil during decay.     

Relationship between plant nitrogen conservation strategies and the dynamics of soil nitrogen in the arid Patagonian Monte, Argentina

During three consecutive years with contrasting precipitation, we analysed the relationship between strategies of N conservation in the dominant plant functional groups (perennial grasses and evergreen shrubs) of the Patagonian Monte and the main components of N cycling in soil. We hypothesised that the different patterns of N conservation in perennial grasses and evergreen shrubs would have direct consequences for soil-N, inorganic-N release and microbial-N flush in soil. In autumn and late spring of 1999, 2000, and 2001, we assessed N and C concentration in green and senesced leaves, N-resorption efficiency and C/N ratio in senesced leaves of three dominant species of each plant functional group. In the soil associated with species of each plant functional group, we determined N and C concentration, potential-N mineralisation, and the associated microbial-N flush. Slow-growing evergreen shrubs exhibited low N-concentration in green leaves, high N-concentration in senesced leaves and low N-resorption from senescing leaves. In contrast, fast-growing perennial grasses showed high N-concentration in green leaves, low N-concentration in senesced leaves, and high N-resorption from senescing leaves. In evergreen shrubs, the maintenance of long-lasting green leaves with low N-concentration was the most important mechanism of N conservation. In contrast, perennial grasses conserved N through high N-resorption from senescing leaves. Soil-N concentration, potential N-mineralisation, and microbial-N flush in the soil were higher underneath evergreen shrubs than beneath perennial grasses. Observed differences, however, were lower than expected considering the quality of the organic matter supplied by each plant fuctional group to the soil. A possible reason for this relatively weak trend may be the capacity of evergreen shrubs to slow down N cycling through low leaf turnover and the presence of secondary compounds in leaves. Alternatively or simultaneously, the weak relationship between plant and soil N could result from shrubs being able to colonise N-poor soils while grasses may preferably occupy fertile microsites previously influenced by the decomposition pathway of evergreen shrubs. Differences between evergreen shrubs and perennial grasses in the mechanisms of plant N-conservation and in components of N cycling in the underlying soil were consistent over the three years of the study with differing precipitation. Inter-annual differences in N concentration in green leaves and in the microbial-N flush in soil indicate that during the wettest year fast-growing perennial grasses would outcompete slow-growing evergreen shrubs and microorganisms for N uptake.     

Soluble phenolics extracted from Larrea divaricata leaves modulate soil microbial activity and perennial grass establishment in arid ecosystems of the Patagonian Monte,Argentina

Plant Ecology - Sheep grazing induces the reduction of perennial grass cover and the increase of shrub cover with high concentration of chemical defences. We analysed the effects of secondary...     

Leafing patterns and leaf traits of four evergreen shrubs in the Patagonian Monte, Argentina

We assessed leafing patterns (rate, timing, and duration of leafing) and leaf traits (leaf longevity, leaf mass per area and leaf-chemistry) in four co-occurring evergreen shrubs of the genus Larrea and Chuquiraga (each having two species) in the arid Patagonian Monte of Argentina. We asked whether species with leaves well-defended against water shortage (high LMA, leaf longevity, and lignin concentration, and low N concentration) have lower leaf production, duration of the leafing period, and inter-annual variation of leafing than species with the opposite traits. We observed two distinctive leafing patterns each related to one genus. Chuquiraga species produced new leaves concentrated in a massive short leafing event (5–48 days) while new leaves of Larrea species emerged gradually (128–258 days). Observed leafing patterns were consistent with simultaneous and successive leafing types previously described for woody plants. The peak of leaf production occurred earlier in Chuquiraga species (mid September) than in Larrea species (mid October–late November). Moreover, Chuquiraga species displayed leaves with the longest leaf lifespan, while leaves of Larrea species had the lowest LMA and the highest N and soluble phenolics concentrations. We also observed that only the leaf production of Larrea species increased in humid years. We concluded that co-occurring evergreen species in the Patagonian Monte displayed different leafing patterns, which were associated with some relevant leaf traits acting as plant defenses against water stress and herbivores. Differences in leafing patterns could provide evidence of ecological differentiation among coexisting species of the same life form.     

Diversity of phenolic compounds and plant traits in coexisting Patagonian desert shrub species of Argentina

Desert shrubs often accumulate different types of phenolic compounds but what determines the amount and diversity of these compounds is an issue scarcely explored. The aim of this study was to assess differences in the amount and diversity of phenolic compounds in leaves among coexisting shrub species differing in rooting depth and leaf turnover. We hypothesized that the diversity and amount of phenolic compounds in leaves of desert shrubs are related to access to soil water through rooting depth, and to leaf turnover. The study was carried out in the Patagonian Monte of Argentina. We collected green leaves of six species representing the dominant shrub morphotypes (tall evergreen, tall deciduous, and medium evergreen shrubs) and assessed lignin concentration and groups of soluble phenols obtained by sequential extraction with ethyl ether, ethyl acetate, and amyl alcohol. We also assessed nitrogen concentration in leaves and leaf mass per unit area (LMA) as traits related to leaf lifespan. The diversity of phenolic compounds was higher in green leaves of tall shrubs with deep rooting depth than in those of medium evergreen shrubs with shallow rooting depth. Diversity of phenolic compounds in green leaves was negatively related to lignin concentration. Evergreen shrubs had higher amount of phenolic compounds in green leaves than deciduous ones and the total amount of phenolic compounds in green leaves was positively related to LMA. We concluded that access to soil water sources and leaf turnover were related to the amount and diversity of phenolic compounds in green leaves of desert shrub species and these results are consistent with those predicted by the resource availability theory for plants from resource-rich and resource-poor habitats.     

Production and turnover rates of shallow fine roots in rangelands of the Patagonian Monte, Argentina

Selective sheep grazing in arid rangelands induces a decrease in total cover and grass cover and an increase in the dominance of shrubs. Both life forms differ in aboveground and belowground traits. We hypothesized that grazing disturbance leads to the replacement of grass by shrub fine roots in the upper soil, and this is reflected in changes in the seasonal dynamics of shallow fine roots at the community level. In two sites representative of non-grazed and grazed vegetation states in the Patagonian Monte, we assessed the canopy structure, and the fine root biomass, N concentration, production, and turnover during two consecutive years. The non-grazed site exhibited higher total, grass, and shrub cover than the grazed site. The grazed site had larger or equal fine root biomass than the non-grazed site except for late spring of the second year. This could be associated with the ability of shrubs to develop dimorphic-root systems occupying the soil freed by grasses at the grazed site, and with the larger contribution of grass than shrub fine roots in relation to an extraordinary precipitation event at the non-grazed site. This was consistent with the N concentration in fine roots. Fine root production was positively correlated to temperature at the grazed site and with precipitation at the non-grazed site. Fine root turnover did not differ between sites. Our results indicate that grazing leads to a shifting in the seasonality and main climatic controls of fine root production, while fine root turnover is mostly affected by changes in soil water conditions.     

Soil Nitrogen in Relation to Quality and Decomposability of Plant Litter in the Patagonian Monte,Argentina

In two consecutive years, we analysed the effect of litter quality, quantity and decomposability on soil N at three characteristic sites of the Patagonian Monte. We assessed (i) concentrations of N, C, lignin and total phenolics and the C/N ratio in senesced leaves as indicators of litter quality of three species of each dominant plant life form (evergreen shrubs and perennial grasses), and (ii) N, and organic-C concentrations, potential N-mineralisation and microbial-N flush in the soil beneath each species. Rate constants of potential decomposition of senesced leaves and N content in decaying leaves during the incubation period were assessed in composite samples of the three sites as indicators of litter decomposability. Further, we estimated for each species leaf-litter production, leaf-litter on soil, and the mass of standing senesced leaves during the senescence period. Senesced leaves of evergreen shrubs showed higher decomposability than those of perennial grasses. Leaf-litter production, leaf-litter on soil, and the mass of standing senesced leaves differed significantly among species. The largest variations in leaf-litter production and leaf-litter on soil were observed in evergreen shrubs. The mass of standing senesced leaves was larger in perennial grasses than in evergreen shrubs. Nitrogen, organic C and potential N-mineralisation in soil were higher underneath evergreen shrubs than beneath perennial grasses, while no significant differences were found in microbial-N flush among life forms. The initial concentrations of C, N and total phenolics of senesced leaves explained together 78% of the total variance observed in the dry mass loss of decaying leaves. Litter decomposition rates explained 98%, 98%, 73%, and 67% of the total variance of soil N, organic C, net-N mineralisation, and microbial-N flush, respectively. We concluded that leaf-litter decomposition rates along with leaf-litter production are meaningful indicators of plant local effects on soil N dynamics in shrublands of the Patagonian Monte, and probably in other similar ecosystem of the world dominated by slow growing species that accumulate a wide variety of secondary metabolites including phenolics. Indicators such as C/N or lignin concentration usually used to predict litter decomposability or local plant effects may not be adequate in the case of slow growing species that accumulate a wide range of secondary metabolites or have long leaf lifespan and low leaf-litter production.     

Mycorrhizal associations in the Patagonian steppe,Argentina

The mycorrhizal status of plant species in north-west Patagonia was examined. Communities representative of Patagonian steppe and marshes were compared with respect to the mycorrhizal status of their species.Most of both native and exotic plant species sampled were arbuscular mycorrhizal (AM). The percentage of species with mycorrhizal association was higher for perennial herbs and shrubs than for annual herbs. The higher ratio of mycorrhizal/nonmycorrhizal (NM) species found for dicotyledons than for monocotyledons, could reflect the presence of a considerable number of NM monocotyledons in the marsh. The mycorrhizal status of plants differed slightly between the steppe and the marsh. In the steppe, native AM species were more frequent than in the marsh. In contrast, in the marsh, the NM species were proportionally more represented than in the steppe. The Juncaceae and Cyperaceae, which include hydrohytic NM plants, accounted for many of these differences. Moreover, the dominant species in the marshes, Juncus arcticus, is a NM species.In the present study, most of species belonging to the same taxonomic family tended to have the same mycorrhizal associations, in agreement with studies on plants from other regions. Exceptions to this general behaviour were observed in the families Cyperaceae, Scrophulariaceae, Berberidaceae and Amaryllidaceae. The most represented families in which mycorrhizal behaviour differed between species of the same family were Asteraceae, Fabaceae and Poaceae. Senecio neaei (Asteraceae) and Boopis australis (Calyceraceae) showed facultative mycorrhizal behaviour.     

Spatial patterns of the germinable soil seed bank of coexisting perennial-grass species in grazed shrublands of the Patagonian Monte

We compared the size and spatial pattern of the germinable soil seed bank (GSB) of the three dominant perennial-grass species in the arid Patagonian Monte of Argentina. These species differ in plant functional traits, seed morphology, and in their preference by grazers. During three consecutive years, we sampled the soil seed bank at 1 m intervals and the intercepted diameter of perennial-grass species and shrub patches along 50-m linear transects at three sites with different grazing pressure. Soil samples were incubated at field capacity during 12 weeks and emerged seedlings counted. The largest-seeded and most xerophytic Stipa speciosa did not form a GSB. The other two grass species with more mesophytic traits and smaller seeds than S. speciosa formed different GSB. Poa ligularis formed a smaller and more heterogeneous GSB than Stipa tenuis. Grazing had not effect on GSB size of P. ligularis but increased the proportion of seeds under dense shrub canopies. In contrast, grazing reduced the size and also increased the proportion of the GSB of S. tenuis under shrub canopies. We conclude that the size and the spatial pattern of the GSB of the dominant perennial-grass species in the arid Patagonian Monte were related not only to seed and plant traits but also to the spatial distribution of grass plants and their exposure to grazers.     

Spatial and temporal litterfall heterogeneity generated by woody species in the Central Monte desert

In arid and semiarid environments, the presence of woody species generates a series of environmental gradients that increase spatial heterogeneity and modify the pattern of distribution of the other species. We postulate that the temporal and spatial variability in litter input generated by woody species is a relevant factor in the generation of edaphic heterogeneity by redistribution of nutrients and the physical effects of litter. The objective of this study was to determine the temporal and spatial variability in the amount of litter input under the canopy of dominant woody plants (Prosopis flexuosa and Larrea divaricata) and in exposed areas at the Ñacuñán Reserve, in the central zone of the Monte desert. Litterfall was collected during 2 years from 30-cm-diameter litter traps distributed at three microsites: under P. flexuosa canopy, under L. divaricata canopy, and in exposed areas. Microhabitats beneath Prosopis showed the highest litter input per m² (between 320 and 527 g/m²), and, consequently, more than 50% of it fell to the soil beneath the canopy of P. flexuosa. Only 10% fell on exposed areas, which exhibited an annual input rate per m² of a lower order of magnitude than the sites under Prosopis. Litterfall presented a peak in summer as a consequence of convective storms, and a second one in autumn due to phenological shedding. Our results suggest that woody species have a central importance in the dynamics of nutrients in arid lands by both the increase of total productivity and litterfall, and the spatial and temporal regulation of litter input.     

A new anisakid species parasitizing the imperial cormorant Phalacrocorax atriceps from the North Patagonian Coast, Argentina

The anisakid species Contracaecum chubutensis n. sp. was found parasitizing the imperial cormorant Phalacrocorax atriceps at 2 different localities, Bahía Bustamante (45 degrees 11'S, 66 degrees 30'W) and Puerto Madryn (42 degrees 47'S, 65 degrees 02'W) on the Patagonian coast. Morphometrical analysis and further studies of adult specimens of C. chubutensis n. sp. and fourth-stage larvae of Contracaecum sp. were done using light and scanning electron microscopy. The possession and disposition of 7 pairs of tail papillae differentiate this new species from C. osculatum, C. radiatum, C. plagiaticium, and C. mirounga. Double-sized spicules separate it from C. variegatum and C. pelagicum. Moreover, the paracloacal papillae disposition and tail shape differentiate this new species from C. travassosi. The presence of bifurcated interlabia separate it from C. ogmorhini and C. margolisi. Three well-marked lip notches, a sharp tail shape, and the of the transversal paracloacal papillae disposition distinguishes C. chubutensis n. sp. from C. magnipapillatum and C. septentrionale. The species here described can be differentiated from C. caballeroi by its bifurcated interlabia, longer spicules, and for possessing more precloacal papillae pairs. It can be separated from C. rudolphii by a thinner interlabia at the base, deeper lip notches, a sharper tail, the disposition of transversal paracloacal papillae, and greater distance between both papillae of the first subventral distal papillae pair. In addition, fourth-stage larvae recovered from the same host and localities were identified and described as Contracaecum sp. Prevalence of adult parasites was 66.7 %, mean intensity was 4.0, and mean abundance was 7.1.     

Seed longevity and fire: germination responses of an exotic perennial herb in NW Patagonian grasslands (Argentina)

Fire affects grassland composition by selectively influencing recruitment. Some exotic species can increase their abundance as a consequence of fire-stimulated seed germination, but response may depend on seed age. Rumex acetosella L. (Polygonaceae, sheep's sorrel) is a cosmopolitan herb that has invaded NW Patagonia's grasslands. This species forms persistent soil seed banks and increases after disturbances, particularly fire. We studied how fire and seed longevity influence R. acetosella germination. In 2008, we conducted laboratory experiments where we exposed different-aged seeds (up to 19 years old) to heat, smoke, charcoal, ash and control treatments. Total percentage germination and mean germination time depended on both seed age and fire treatment. Germination of younger seeds decreased with increasing temperature. There was no general pattern in germination responses of different-aged seeds to smoke, charcoal and ash. While smoke improved the germination of fresh seeds, charcoal decreased germination. Germination of untreated seeds was negatively correlated with seed age, and mean germination time increased with seed age. In most treatments, fresh seeds had lower germination than 1-5-year-old seeds, indicating an after-ripening requirement. Smoke stimulates R. acetosella germination, causing successful recruitment during post-fire conditions. Fresh seeds are particularly responsive to fire factors, possibly because they have not experienced physical degradation and are more receptive to environmental stimuli. Knowing the colonisation potential from the soil seed bank of this species during post-fire conditions will allow us to predict their impact on native communities.     

Assessment of habitat quality for four small mammal species of the Monte Desert, Argentina

In the temperate desert of Argentina, the combined action of climatic and anthropogenic factors has contributed to the formation of a highly heterogeneous landscape. In the central region of the Monte desert, four small mammal species (Eligmodontia typus, Calomys musculinus, Akodon molinae and Graomys griseoflavus) coexist and show different habitat uses in response to spatial variability. Three main habitat types are present in the region: mesquite forest, the creosotebush community and sand dunes. These habitat types are present also in the surrounding grazing area.The objective of this study was to determine habitat quality for these species in a protected area (Reserve MaB Ñacuñán) and in the adjacent grazed area. For each species we estimated demographic parameters that are highly correlated to fitness in each habitat, and for both treatments (protected and grazed).We found that the protected area offered a higher quality habitat than the grazed area for all species, but principally A. molinae and G. griseoflavus. At a local scale, we found that A. molinae and C. musculinus clearly showed higher fitness in the more complex habitats as the creosotebush community and the mesquite forest. In contrast, for E. typus, open and simplest patches, such as sand dunes, were optimal for its survival and reproduction.     

Multi-scale patterns of habitat use by wild boar in the Monte Desert of Argentina

A large number of protected areas worldwide have been impacted by biological invasions, threatening the biodiversity they aim to protect. The wild boar (Sus scrofa) is one of the most threatening invasive species in Argentina, already occupying many ecoregions, including the central Monte Desert. However, there are no studies regarding the use that wild boars make of this invaded biome and what factors (climate or landscape) determine or contribute to the establishment of this species. The objectives of this study were to assess habitat use of the wild boar at spatial and temporal scales in the central Monte Desert, and to assess if climatic factors influence its abundance. Our results show that, at habitat-level the wild boar exhibited preferences for a particular habitat (Larrea shrubland) for feeding. At microhabitat-level, we found a positive association between herb cover and wild boar presence. In addition, we found a strong and positive association between the number of days with low temperatures and the number of wild boar signs registered. Therefore, we consider that in the central Monte Desert, habitat selection by wild boars is most likely determined by a maximization of food intake and a minimization of exposure to high temperature.     

A new Huffmanela species (Nematoda: Trichosomoididae) parasitizing atherinid fishes in north Patagonian Gulfs, Argentina

Huffmanela moraveci n. sp. (Nematode: Trichosomoididae) is described from the skin of fins and gills of the silversides Odontesthes smitti Lahille, 1949, and Odontesthes nigricans Richardson, 1848 (Pisces: Atheriniformes), from southwestern Atlantic coast, on the basis of the morphology of adults and eggs found in the host tissues. Huffmanela moraveci n. sp. can be distinguished from Huffmanela carcharini, H. banningi, H. japonica, H. mexicana, H. paronai, H. schouteni, H. shikokuensis, H. branchialis, H. filamentosa, H. ossicola, and H. lata by the size and shape of the eggs. The adult stage of H. moraveci n. sp. can be differentiated from H. hufmani and H. canadensis by the position of the nerve ring and by the total esophagus length/body length ratio. Huffmanela moraveci n. sp. also differs from H. huffmani by the presence of bacillary bands and by the body length. This is the first report of a species of Huffmanela in South America, the first report of a species of Huffmanela with Odontesthes spp. as host, and the third known adult form in the genus.