Luciana Ghermandi

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’. Abbreviation: MRPP = Multi response permutation procedures. Nomenclature: Correa (1969–1999).

Leaf-cutting ant nests near roads increase fitness of exotic plant species in natural protected areas

Understanding the mechanisms that promote the invasion of natural protected areas by exotic plants is a central concern for ecology. We demonstrated that nests of the leaf-cutting ant, Acromyrmex lobicornis, near roadsides promote the abundance, growth and reproduction of two exotic plant species, Carduus nutans and Onopordum acanthium, in a national park in northern Patagonia, Argentina and determine the mechanisms that produce these effects. Refuse dumps (RDs) from ant nests have a higher nutrient content than nearby non-nest soils (NNSs); foliar nutrient content and their 15N isotopic signature strongly suggest that plants reach and use these nutrients. Both species of exotic plants in RDs were 50–600% more abundant; seedlings had 100–1000% more foliar area and root and leaf biomass; and adult plants produced 100–300% more seeds than nearby NNS plants. Plants can thus gain access to and benefit from the nutrient content of ant RD, supporting the hypotheses that enhanced resource availability promotes exotic plant performance that could increase the likelihood of biological invasions. The two exotics produce an estimated of 8 385 000 more seeds ha−1 in areas with ant nests compared with areas without; this exceptional increase in seed production represents a potential threat to nearby non-invaded communities. We propose several management strategies to mitigate this threat. Removal efforts of exotics should be focused on ant RDs, where plants are denser and represent a higher source of propagules.

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

Abstract 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. Nomenclature: Correa (1969–1998).

Ecological engineering by a native leaf-cutting ant increases the performance of exotic plant species

Numerous mechanisms are proposed to explain why exotic plants successfully invade natural communities. However, the positive effects of native engineers on exotic plant species have received less consideration. We tested whether the nutrient-rich soil patches created by a native ecological engineer (refuse dumps from the leaf-cutting ant Acromyrmex lobicornis) increase the performance of exotic more than native plants. In a greenhouse experiment, individuals from several native and exotic species were planted in pots with refuse dumps (RDs) and non-nest soils (NNSs). Total plant biomass and foliar nutrient content were measured at the end of the experiment. We also estimated the cover of exotic and native plant species in external RDs from 54 field ant nests and adjacent areas. Greenhouse plants showed more biomass and foliar nutrient content in RDs than in NNS pots. Nevertheless, differences in the final mean biomass among RD and NNS plants were especially great in exotics. Accordingly, the cover of exotic plants was higher in field RDs than in adjacent, non-nest soils. Our results demonstrated that plants can benefit from the enhanced nutrient content of ant RDs, and that A. lobicornis acts as an ecosystem engineer, creating a substrate that especially increases the performance of exotics. This supports the fluctuating resource hypothesis as a mechanism to promote biological invasions, and illustrates how this hypothesis may operate in nature. Since ant nests and exotic plants are more common in disturbed than in pristine environments, the role of ant nests in promoting biological invasions might be of particular interest. Proposals including the use of engineer species to restore disturbed habitats should be planned with caution because of their potential role in promoting invasions.

Effect of fire on recruitment of two dominant perennial grasses with different palatability from semi-arid grasslands of NW Patagonia (Argentina)

Fire is a non-selective disturbance that impacts equally plant species that could be selected differentially by livestock. Post-fire recruitment dynamics is an important ecological process that has been barely studied in Patagonian grass species. This work analyzes the effect of fire on seed germination, seedling growth, and survival of Pappostipa speciosa (ex Stipa speciosa) and Festuca pallescens, two dominant perennial grasses from NW Patagonia that differ in palatability. We hypothesized that physical and chemical factors derived from fire differentially affect recruitment of these species. We performed experiments in the field and under laboratory and greenhouse conditions to study the integral effect of fire and of related abiotic factors (i.e., smoke, heat, charcoal, and ash) on different phases of recruitment of both species. Experimental burning promoted P. speciosa emergence over time, but they did not affect F. pallescens total emergence. Experimental burning decreased P. speciosa seedling growth (i.e., few leaves and small size), but they did not affect seedling survival. Smoke from laboratory experiments stimulated P. speciosa germination. Exposing F. pallescens seeds to 120°C decreased germination and seedling growth. Fire might act as a selective force on recruitment of both species, as well as changing competitive interactions during postfire regeneration. The effect of fire on the recruitment dynamics of the studied species depended strongly on both intrinsic species characteristics and meteorological conditions.

Effects of fire severity on early recovery of Patagonian steppes

Fire severity influences the recovery and biodiversity of plant communities. We compared the early post-fire recovery of plant functional groups in two sites (one burnt with high severity and one with low severity) in the north-western Patagonian steppe. We sampled the post-fire soil seed banks, as well as the composition and cover of aboveground plant species, and calculated richness, diversity and the percentage of exotic species. Low fire severity increased the diversity of aboveground vegetation and seed banks, whereas high fire severity reduced the cover of all functional groups, with the exception of fugitive species. Tiller production of perennial grasses decreased due to bud bank mortality. In the seed bank, high severity fire reduced fugitive species and increased exotic species. Fugitive species need fire to recruit, and the combination of fire cues and post-fire environmental conditions can explain the high abundance of these species after fire. In Patagonia, the use of fire is not considered a management option because fires are believed to be dangerous to humans, goods and services. However, north-western Patagonia is sparsely populated and the frequent wildfires pose no danger. It is necessary to increase the knowledge of the ecological processes of Patagonian ecosystems related to fire regime and land use, to achieve a reasonable balance between sustainable use and conservation in this region.

Growth and reproductive post-fire responses of two shrubs in semiarid Patagonian grasslands

Variation in fire intensity affects the post-fire survival and growth of shrubs. We examined effects of maximum fire temperature (a proxy for fire intensity) on the survival, growth and reproductive performance of the shrubs Mulinum spinosum and Senecio bracteolatus at 1 and 2 years post-fire in north-west Patagonian grasslands. We applied two fire temperature treatments to plants of each species. All M. spinosum plants survived fire treatments but high fire temperature notably decreased survival of S. bracteolatus. Bud position and plant architecture probably influenced shrub survival. During the first growing season, M. spinosum did not produce seeds whereas S. bracteolatus plants burned at low temperature produced three times more seeds and bigger seeds than unburned plants. Also, seeds from burned plants of S. bracteolatus had higher germinability than seed from unburned plants. High survival and resprouting capacity of M. spinosum even after high fire intensity indicate that this species might be less affected by changes in fire regime.

Fire as a stimulant of shrub recruitment in northwestern Patagonian (Argentina) grasslands

One strategy of plant survival during post-fire succession is to persist and regenerate by recruiting new individuals from a fire-resistant seed bank. The heat, smoke, and charcoal released during plant combustion may act (individually or in combination) as a cue for post-fire seed germination. Fabiana imbricata is a shrub that forms persistent seed banks in the northwestern Patagonian grasslands and shows a high recruitment from seeds during post-fire succession. Mathematical models showed that this species is advancing over the grasslands in response to fires. To corroborate these findings, we studied the role of fire on F. imbricata seed germination. In order to achieve this, a factorial experiment was designed in laboratory conditions to study the effect of heat, charcoal, smoke, scarification, and their interactions on F. imbricata seed germination. Seeds treated with the higher temperatures required a longer period of time to germinate, thus, significantly affecting the mean germination time. Total germination percentages in F. imbricata were significantly enhanced by smoke and scarification, but the interaction of heat, smoke, and scarification was more important than the effect of each fire factor alone. The positive response to fire cues exhibited by F. imbricata indicates that this species would have an adaptive advantage to colonize these grasslands if fire frequency increased, as predicted for this environment. Hence, fire will contribute to the grassland encroachment by this species and, therefore, to the loss of biodiversity and productivity of northwestern Patagonian grasslands.

From leaves to landscape: A multiscale approach to assess fire hazard in wildland-urban interface areas.

The overlapping zone between urbanization and wildland vegetation, known as the wildland urban interface (WUI), is often at high risk of wildfire. Human activities increase the likelihood of wildfires, which can have disastrous consequences for property and land use, and can pose a serious threat to lives. Fire hazard assessments depend strongly on the spatial scale of analysis. We assessed the fire hazard in a WUI area of a Patagonian city by working at three scales: landscape, community and species. Fire is a complex phenomenon, so we used a large number of variables that correlate a priori with the fire hazard. Consequently, we analyzed environmental variables together with fuel load and leaf flammability variables and integrated all the information in a fire hazard map with four fire hazard categories. The Nothofagus dombeyi forest had the highest fire hazard while grasslands had the lowest. Our work highlights the vulnerability of the wildland-urban interface to fire in this region and our suggested methodology could be applied in other wildland-urban interface areas. Particularly in high hazard areas, our work could help in spatial delimitation policies, urban planning and development of plans for the protection of human lives and assets.

Fire regime from 1973 to 2011 in north-western Patagonian grasslands

Fire is one of the most important disturbances in terrestrial ecosystems and has major ecological and socioeconomic impacts. Fire regime describes the variation of individual fire events in time and space. Few studies have characterised the fire regime in grasslands in spite of the importance of these ecosystems. The aim of this study was to describe the recent fire regime (from 1973 to 2011) of north-western Patagonian grasslands in terms of seasonality, frequency and burned area. Our study area covered 560 000 ha and we used a remote sensing approach combined with statistics obtained from operational databases. Fires occur during the summer in 2 of every 3 years with a frequency of 2.7 fires per year and a mean size of 823 ha. Fire size distribution is characterised by many small fires and few large ones which would respond to a distribution from the power law family. Eighty per cent of the total area affected by fire was burned in the span of a few years, which were also widespread fire years in forests and woodlands of north-western Patagonia. This work contributes to general knowledge about fire regimes in grasslands and we expect that our results will serve as a reference to further fire regime research.