No Sul do Brasil ocorrem mosaicos naturais de vegetação campestre e florestal, sob um clima que é favorável às formações florestais. O processo de expansão florestal tem sido observado principalmente sobre áreas de campo excluído de perturbações de fogo e pastejo. Estudos sobre padrões de vegetação lenhosa associados a variáveis ambientais locais em áreas de campo sujeitas à expansão florestal constituem uma importante ferramenta para avaliação da dinâmica da vegetação considerando mudanças ambientais ou de regime de perturbação. No intuito de inferir sobre os processos relacionados à expansão florestal sobre campos no Sul do Brasil, este estudo busca os padrões atuais na composição e estrutura da vegetação lenhosa sobre uma área de campo excluído de manejo há 34 anos e as possíveis associações destes com variáveis espaciais e ambientais. Para tanto, amostramos a vegetação lenhosa em 110 unidades amostrais circulares (6m de raio) estabelecidas de forma sistemática (equidistantes 100m) sobre áreas originalmente mapeadas, em 1985, como vegetação campestre ou vassoural. A amostragem foi estratificada para o critério de inclusão baseado no diâmetro do caule a altura do solo (DAS). As unidades amostrais de 6m de raio contemplavam os indivíduos com DAS5cm e estas incluíram três sub-parcelas de 1m de raio para amostragem proporcional de indivíduos menores (DAS<5cm, mas com altura superior a 1m). Como variáveis ambientais, a distância mínima de uma área-fonte florestal e fatores químicos e físicos do solo foram avaliados para cada unidade amostral. Os dados de vegetação foram analisados através de parâmetros fitossociológicos, sendo os padrões, e suas relações com o ambiente, acessados por análise exploratória conjugando técnicas estatísticas uni e multivariadas. Os resultados revelam que em áreas campestres excluídas de fogo e pastejo, como a que foi estudada, ocorre um processo espacial e temporal relacionado com a distância da área fonte florestal, mostrando um gradiente sucessional. Quanto mais próximo da antiga borda florestal, as comunidades de espécies lenhosas são mais antigas e caracterizadas por espécies mais tipicamente florestais. A medida que o processo se afasta da borda encontram-se comunidades em estágios sucessionais intermediários, onde os arbustos campestres gradualmente cedem lugar a espécies florestais pioneiras, nucleadoras de vegetação florestal, as quais atraem dispersores e criam condições favoráveis ao estabelecimento de mais espécies florestais. E nas áreas mais distantes da borda ainda predominam arbustos campestres (e.g. Baccharis uncinella). Eles gradativamente suprimem o estrato herbáceo de gramíneas pela sua abundância e, em alguma medida, facilitam o estabelecimento das espécies florestais pioneiras. Nesses processos, o solo se modifica em direção a condições de maior acúmulo de matéria orgânica e de maior fertilidade, mais favoráveis ao estabelecimento de espécies florestais. / In South of Brazil there are forest-grassland mosaics under a climate suit for forest development. The forest expansion process has been observed mainly over grasslands excluded from cattle grazing and fire disturbances. Studies accessing patterns of woody vegetation linked to local environmental variables in grasslands areas subject to forest expansion constitute an important tool for vegetation dynamics evaluation considering environmental or disturbance management changes. Our purpose is access the current woody vegetation patterns and their potential linking with spatial and environmental variables on a grassland area excluded from management since 34 years ago. For this we surveyed woody species on 110 sampling units systematically plotted (halfway 100m) over areas that were originally mapped at 1985 as grasslands or shrublands. The sampling was stratified according to the inclusion criterion based on the stem diameter at the height of the soil (DHS). In the 6m ray sampling units all individuals with DHS5cm were surveyed and, inside them, three sub-plots of 1m of ray were used to sample proportional smaller individuals (DHS<5cm, but with more than 1m height). As environmental variables, the minimum distance of a forest area-source, and chemical and physical soil characteristics were considered for each sampling unit. The vegetation data and patterns were analyzed through phytosociological parameters and by exploratory analysis conjugating united and multivariate statistical techniques. The results revealed that grassland areas excluded of fire and grazing, as the one that it was studied, are suitable to spatial and temporal processes related to the distance of the source-forest area, showing a sucessional gradient. Close to the old forest border, the woody species communities are older and characterized by typical forest species. As the process stands back of the border the communities are in intermediate sucessional stage, where grassland shrubs gradually give up place to pioneer forest species, nurse species, which attract dispersers and create favorable conditions to the establishment of more forest species, allowing forest expansion process. In the most distant border areas grassland shrubs still prevail (e.g. Baccharis uncinella). They gradually suppress the grasses stratum because of their abundance and, in some way, facilitate the establishment of pioneer forest species. In those processes, the soil modifies towards conditions of larger accumulation of organic matter and larger fertility, more favorable to the establishment of forest species.

No Sul do Brasil ocorrem mosaicos naturais de vegetação campestre e florestal, sob um clima que é favorável às formações florestais. O processo de expansão florestal tem sido observado principalmente sobre áreas de campo excluído de perturbações de fogo e pastejo. Estudos sobre padrões de vegetação lenhosa associados a variáveis ambientais locais em áreas de campo sujeitas à expansão florestal constituem uma importante ferramenta para avaliação da dinâmica da vegetação considerando mudanças ambientais ou de regime de perturbação. No intuito de inferir sobre os processos relacionados à expansão florestal sobre campos no Sul do Brasil, este estudo busca os padrões atuais na composição e estrutura da vegetação lenhosa sobre uma área de campo excluído de manejo há 34 anos e as possíveis associações destes com variáveis espaciais e ambientais. Para tanto, amostramos a vegetação lenhosa em 110 unidades amostrais circulares (6m de raio) estabelecidas de forma sistemática (equidistantes 100m) sobre áreas originalmente mapeadas, em 1985, como vegetação campestre ou vassoural. A amostragem foi estratificada para o critério de inclusão baseado no diâmetro do caule a altura do solo (DAS). As unidades amostrais de 6m de raio contemplavam os indivíduos com DAS5cm e estas incluíram três sub-parcelas de 1m de raio para amostragem proporcional de indivíduos menores (DAS<5cm, mas com altura superior a 1m). Como variáveis ambientais, a distância mínima de uma área-fonte florestal e fatores químicos e físicos do solo foram avaliados para cada unidade amostral. Os dados de vegetação foram analisados através de parâmetros fitossociológicos, sendo os padrões, e suas relações com o ambiente, acessados por análise exploratória conjugando técnicas estatísticas uni e multivariadas. Os resultados revelam que em áreas campestres excluídas de fogo e pastejo, como a que foi estudada, ocorre um processo espacial e temporal relacionado com a distância da área fonte florestal, mostrando um gradiente sucessional. Quanto mais próximo da antiga borda florestal, as comunidades de espécies lenhosas são mais antigas e caracterizadas por espécies mais tipicamente florestais. A medida que o processo se afasta da borda encontram-se comunidades em estágios sucessionais intermediários, onde os arbustos campestres gradualmente cedem lugar a espécies florestais pioneiras, nucleadoras de vegetação florestal, as quais atraem dispersores e criam condições favoráveis ao estabelecimento de mais espécies florestais. E nas áreas mais distantes da borda ainda predominam arbustos campestres (e.g. Baccharis uncinella). Eles gradativamente suprimem o estrato herbáceo de gramíneas pela sua abundância e, em alguma medida, facilitam o estabelecimento das espécies florestais pioneiras. Nesses processos, o solo se modifica em direção a condições de maior acúmulo de matéria orgânica e de maior fertilidade, mais favoráveis ao estabelecimento de espécies florestais. / In South of Brazil there are forest-grassland mosaics under a climate suit for forest development. The forest expansion process has been observed mainly over grasslands excluded from cattle grazing and fire disturbances. Studies accessing patterns of woody vegetation linked to local environmental variables in grasslands areas subject to forest expansion constitute an important tool for vegetation dynamics evaluation considering environmental or disturbance management changes. Our purpose is access the current woody vegetation patterns and their potential linking with spatial and environmental variables on a grassland area excluded from management since 34 years ago. For this we surveyed woody species on 110 sampling units systematically plotted (halfway 100m) over areas that were originally mapped at 1985 as grasslands or shrublands. The sampling was stratified according to the inclusion criterion based on the stem diameter at the height of the soil (DHS). In the 6m ray sampling units all individuals with DHS5cm were surveyed and, inside them, three sub-plots of 1m of ray were used to sample proportional smaller individuals (DHS<5cm, but with more than 1m height). As environmental variables, the minimum distance of a forest area-source, and chemical and physical soil characteristics were considered for each sampling unit. The vegetation data and patterns were analyzed through phytosociological parameters and by exploratory analysis conjugating united and multivariate statistical techniques. The results revealed that grassland areas excluded of fire and grazing, as the one that it was studied, are suitable to spatial and temporal processes related to the distance of the source-forest area, showing a sucessional gradient. Close to the old forest border, the woody species communities are older and characterized by typical forest species. As the process stands back of the border the communities are in intermediate sucessional stage, where grassland shrubs gradually give up place to pioneer forest species, nurse species, which attract dispersers and create favorable conditions to the establishment of more forest species, allowing forest expansion process. In the most distant border areas grassland shrubs still prevail (e.g. Baccharis uncinella). They gradually suppress the grasses stratum because of their abundance and, in some way, facilitate the establishment of pioneer forest species. In those processes, the soil modifies towards conditions of larger accumulation of organic matter and larger fertility, more favorable to the establishment of forest species.

No Sul do Brasil ocorrem mosaicos naturais de vegetação campestre e florestal, sob um clima que é favorável às formações florestais. O processo de expansão florestal tem sido observado principalmente sobre áreas de campo excluído de perturbações de fogo e pastejo. Estudos sobre padrões de vegetação lenhosa associados a variáveis ambientais locais em áreas de campo sujeitas à expansão florestal constituem uma importante ferramenta para avaliação da dinâmica da vegetação considerando mudanças ambientais ou de regime de perturbação. No intuito de inferir sobre os processos relacionados à expansão florestal sobre campos no Sul do Brasil, este estudo busca os padrões atuais na composição e estrutura da vegetação lenhosa sobre uma área de campo excluído de manejo há 34 anos e as possíveis associações destes com variáveis espaciais e ambientais. Para tanto, amostramos a vegetação lenhosa em 110 unidades amostrais circulares (6m de raio) estabelecidas de forma sistemática (equidistantes 100m) sobre áreas originalmente mapeadas, em 1985, como vegetação campestre ou vassoural. A amostragem foi estratificada para o critério de inclusão baseado no diâmetro do caule a altura do solo (DAS). As unidades amostrais de 6m de raio contemplavam os indivíduos com DAS5cm e estas incluíram três sub-parcelas de 1m de raio para amostragem proporcional de indivíduos menores (DAS<5cm, mas com altura superior a 1m). Como variáveis ambientais, a distância mínima de uma área-fonte florestal e fatores químicos e físicos do solo foram avaliados para cada unidade amostral. Os dados de vegetação foram analisados através de parâmetros fitossociológicos, sendo os padrões, e suas relações com o ambiente, acessados por análise exploratória conjugando técnicas estatísticas uni e multivariadas. Os resultados revelam que em áreas campestres excluídas de fogo e pastejo, como a que foi estudada, ocorre um processo espacial e temporal relacionado com a distância da área fonte florestal, mostrando um gradiente sucessional. Quanto mais próximo da antiga borda florestal, as comunidades de espécies lenhosas são mais antigas e caracterizadas por espécies mais tipicamente florestais. A medida que o processo se afasta da borda encontram-se comunidades em estágios sucessionais intermediários, onde os arbustos campestres gradualmente cedem lugar a espécies florestais pioneiras, nucleadoras de vegetação florestal, as quais atraem dispersores e criam condições favoráveis ao estabelecimento de mais espécies florestais. E nas áreas mais distantes da borda ainda predominam arbustos campestres (e.g. Baccharis uncinella). Eles gradativamente suprimem o estrato herbáceo de gramíneas pela sua abundância e, em alguma medida, facilitam o estabelecimento das espécies florestais pioneiras. Nesses processos, o solo se modifica em direção a condições de maior acúmulo de matéria orgânica e de maior fertilidade, mais favoráveis ao estabelecimento de espécies florestais. / In South of Brazil there are forest-grassland mosaics under a climate suit for forest development. The forest expansion process has been observed mainly over grasslands excluded from cattle grazing and fire disturbances. Studies accessing patterns of woody vegetation linked to local environmental variables in grasslands areas subject to forest expansion constitute an important tool for vegetation dynamics evaluation considering environmental or disturbance management changes. Our purpose is access the current woody vegetation patterns and their potential linking with spatial and environmental variables on a grassland area excluded from management since 34 years ago. For this we surveyed woody species on 110 sampling units systematically plotted (halfway 100m) over areas that were originally mapped at 1985 as grasslands or shrublands. The sampling was stratified according to the inclusion criterion based on the stem diameter at the height of the soil (DHS). In the 6m ray sampling units all individuals with DHS5cm were surveyed and, inside them, three sub-plots of 1m of ray were used to sample proportional smaller individuals (DHS<5cm, but with more than 1m height). As environmental variables, the minimum distance of a forest area-source, and chemical and physical soil characteristics were considered for each sampling unit. The vegetation data and patterns were analyzed through phytosociological parameters and by exploratory analysis conjugating united and multivariate statistical techniques. The results revealed that grassland areas excluded of fire and grazing, as the one that it was studied, are suitable to spatial and temporal processes related to the distance of the source-forest area, showing a sucessional gradient. Close to the old forest border, the woody species communities are older and characterized by typical forest species. As the process stands back of the border the communities are in intermediate sucessional stage, where grassland shrubs gradually give up place to pioneer forest species, nurse species, which attract dispersers and create favorable conditions to the establishment of more forest species, allowing forest expansion process. In the most distant border areas grassland shrubs still prevail (e.g. Baccharis uncinella). They gradually suppress the grasses stratum because of their abundance and, in some way, facilitate the establishment of pioneer forest species. In those processes, the soil modifies towards conditions of larger accumulation of organic matter and larger fertility, more favorable to the establishment of forest species.

The origin and classification of open and mosaic ecosystems, particularly in the tropics and subtropics, have led to controversy worldwide. This has affected biodiversity conservation and, in some cases, promoted the establishment of afforestation projects based on the assumption that open and mosaic ecosystems are degraded forests. Although this initiative can have benefits in terms of carbon storage and climate mitigation if carefully planned and managed, it can also cause biodiversity loss and degradation when afforestation takes place in areas that were previously open ecosystems, or where unsuitable species are used. Madagascar, a world biodiversity hotspot, is one of the countries targeted for the implementation of afforestation projects. The Central Highlands of Madagascar, dominated by grassland matrix with forest patches, is the main region targeted. The nature and origin of the landscape are hotly debated, however, and it is not clear whether these open ecosystems are ancient or anthropogenically derived. Understanding of landscape history is therefore required to identify and conserve ancient open ecosystems, and to distinguish them from areas that have been deforested by people. This research aims to reconstruct the vegetation history and environmental change in the Central Highlands of Madagascar during the Holocene using palaeoecological methods, in order to inform appropriate conservation and management plans. We provide new records of vegetation, hydrological change, fire and herbivory activities by using a multiproxy approach, which includes fossil pollen, stable carbon isotopes, diatoms, charcoal and coprophilous spores, that allows for a comprehensive investigation into the history and drivers of vegetation change. Sediment cores were collected from two sites, Tampoketsa-Ankazobe wetland and Lake Dangovavy, located in the eastern and western slopes of the highlands, respectively. Results indicated that the surrounding area of both sites was composed of mosaic ecosystems, comprising of forest patches of variable extent in a matrix of open grassland and ericoid shrubland vegetation, at least from the Early and Mid-Holocene to ca. 1000 cal years BP, driven mainly by climate variability and fire occurrence. In Tampoketsa-Ankazobe wetland (eastern slopes), the vegetation was characterised by a mosaic of ericoid shrubland and mid-elevation forest taxa, between ca. 11 200 and 8300 cal years BP, under warm/wet period and low fire occurrence. The vegetation in the area changed to a mosaic of ericoid shrubland with more dominance of high-elevation forest from ca. 8300 to 1000 cal years BP under a drier climate and consistent low fire occurrence. The abundance of shrubs and trees during those two periods were confirmed by the dominance of C3 plants as reflected by the stable carbon isotopes results, and coincided with low herbivory activities in the TampoketsaAnkazobe site from ca. 11 200 to 1000 cal years BP. In parallel, the pollen record from Lake Dangovavy (western slopes), between ca. 6200 and 5400 cal years BP, suggests a mosaic ecosystem, dominated by more C3 montane grass, ericoid shrubland and high elevation forest patches promoted by cool/dry climate with low fire occurrence and herbivory activities. Between ca. 5400 and 4200 cal years BP, vegetation in the area was dominated by a mosaic of ericoid shrubland and mid-elevation forest under a wetter period, moderate fire occurrence, and herbivory activities. This mosaic was controlled by climate, fire refugia and herbivory feedbacks. The vegetation changed into a forest-savanna mosaic with an abundance of grassland and pioneer/fire-resistant trees between ca. 4200 and 3000 cal years BP. The period was characterised by an initial increase of local fire followed by a regional drought event. This suggests that a threshold might have been reached, with a resulting shift in vegetation composition. Between ca. 3000 and 1000 cal years BP, reoccurrence of ericoid shrubland with woodland savanna taxa was recorded in the area. The vegetation was conditioned by variation of climate from wet (until ca. 2000 cal years BP) to dry period (ca. 2000–⁠1000 cal years BP) with moderate fire occurrence and herbivory activities. In addition, stable carbon isotope results show that between ca. 6200 to 1000 cal years BP, the site was characterised by C3 plants. During the last ca.1000 cal years BP, pollen records from both sites in the Central Highlands of Madagascar showed a shift to a more open landscape dominated by grassland. Trees and shrubland in the highlands experienced a massive decrease and this correlated with an abundance of C4 plants associated with reduced diversity. The shift of vegetation during this period was likely a result of a centennial severe drought period at ca. 950 cal years BP, as recorded in the literature and confirmed by the peak in aerophilous taxa in our diatom record. The drought was followed by a dramatic increase of fire occurrence and herbivory activities in the region, as recorded in the charcoal and spore records from both sites, indicating human activities at ca. 700 and 500 cal years BP for Lake Dangovavy and Tampoketsa-Ankazobe wetland, respectively. Though the vegetation at both sites in Central Highlands of Madagascar was very dynamic until ca. 1000 cal years BP, complex interactions between climate and fire allowed the forest and ericoid elements to persist, consistent with a heterogeneous mosaic landscape. This changed from 1000 years ago with the occurrence of a regional severe drought event followed by an increase in human activities leading to an increase of grass, a decline in forest and ericoid elements. Our findings suggest that although, the eastern and western slopes in Madagascar might have different vegetation histories over time as a response to the complex climatic-fire drivers at least until ca. 1000 cal years BP, they both: a) Contained ancient open ecosystems such as grasslands and/or ericoid shrubland, and a mosaic landscape which should be considered typical of the highland region. b) Experienced a loss of forest, woodland and mosaic elements, a trend that is consistent with the anthropogenic conversion of some forests to grasslands since ca. 1000 cal years BP. Such findings have implications in terms of conservation, fire management and afforestation projects in the Central Highlands, and provide additional knowledge that contributes to the understanding of its ecological processes and history prior to human arrival on the island. Indeed: 1) Ancient grasslands and ericoid shrubland need to be identified and conserved because of their antiquity and unique biodiversity. To date, there has been some focus on ancient grasslands, but the presence of ancient heathlands has not been discussed. 2) It is important to distinguish ancient from derived grasslands and to target the latter for reforestation, using species that are typical of the remaining forest patches. 3) Fire management should be conducted at a local scale and should incorporate the landscape fire history, considering, for example, the differences between two slopes in the Central Highlands.