Caracterización de ecotonos entre biomas de bosque y biomas abiertos en un área protegida en Uruguay

Verónica Etchebarne-Palla; Oscar Blumetto; Ángel M. Segura; Ariel A. Farías; Laura Cappuccio; Álvaro Soutullo

doi:10.25260/EA.22.32.2.0.1801

Autores/as

Verónica Etchebarne-Palla Departamento de Ecología y Gestión Ambiental, Centro Universitario de la Región Este, Universidad de la República. Maldonado, Uruguay. Área Biodiversidad y Conservación, Museo Nacional de Historia Natural, Ministerio de Educación y Cultura. Montevideo, Uruguay
Oscar Blumetto Programa de Producción y Sustentabilidad Ambiental, Estación Experimental INIA Las Brujas, Instituto Nacional de Investigación Agropecuaria. Canelones, Uruguay
Ángel M. Segura Modelización y Análisis de recursos naturales, Centro Universitario Regional Este, Universidad de la República. Rocha, Uruguay
Ariel A. Farías Departamento de Ecología y Gestión Ambiental, Centro Universitario de la Región Este, Universidad de la República. Maldonado, Uruguay. Center of Applied Ecology and Sustainability (CAPES), P. Universidad Catoìlica de Chile. Santiago, Chile
Laura Cappuccio Departamento de Ecología y Gestión Ambiental, Centro Universitario de la Región Este, Universidad de la República. Maldonado, Uruguay
Álvaro Soutullo Departamento de Ecología y Gestión Ambiental, Centro Universitario de la Región Este, Universidad de la República. Maldonado, Uruguay

DOI:

https://doi.org/10.25260/EA.22.32.2.0.1801

Palabras clave:

árboles, arbustos, ganadería, fuego, ovino, vacuno, pastizales, arbustales, conservación

Resumen

La teoría de Estados Alternativos de Biomas (EAB) contribuye a explicar la presencia de biomas abiertos (pastizales, arbustales) en ambientes suficientemente cálidos y húmedos para el desarrollo de biomas de bosque. En estos ambientes se pueden encontrar áreas dominadas por biomas abiertos que, según esta teoría, responden principalmente al fuego y a la herbivoría. El objetivo de este trabajo es identificar si existen diferentes tipos de ecotonos entre biomas de bosque-biomas abiertos en base a la composición leñosa, describirlos y explorar su relación con el manejo ganadero y las condiciones ambientales, utilizando como caso de estudio un área protegida en Uruguay. Muestreamos la vegetación leñosa y otras variables ambientales en transectas (delimitadas en 200 m) en 10 sitios que abarcan cuatro manejos ganaderos. Identificamos dos tipos de ecotonos entre bioma de bosque y bioma abierto. Uno, dominado por arbustal, y otro, dominado por pastizal. Nuestros resultados concuerdan con la teoría de EAB en que los mecanismos que parecen determinar la composición leñosa y la estructura de la vegetación en ecotonos son el ganado y uso de fuego o segadora. Los cambios en la estructura de la vegetación y en la riqueza de leñosas fue, en parte, abrupto, en coincidencia con lo que propone esta teoría. Sin embargo, la composición también interactuó con factores de sitio como la rocosidad y la distancia al curso de agua, lo cual sugiere que variables ambientales a nivel de sitio estarían actuando de forma sinérgica con la remoción de la vegetación para determinar la composición de la vegetación. Encontramos que estos ecotonos son sensibles a manejos y variables ambientales, por lo que es necesario incorporar esta heterogeneidad de paisaje y de sitio para articular la actividad productiva con la conservación de la naturaleza.

Citas

de Abreu, R. C. R., and G. Durigan. 2011. Changes in the plant community of a Brazilian grassland savannah after 22 years of invasion by Pinus elliottii engelm. Plant Ecology and Diversity 4:269-278. https://doi.org/10.1080/17550874.2011.594101.

Altesor, A., M. Oesterheld, E. Leoni, F. Lezama, and C. Rodríguez. 2005. Effect of grazing on community structure and productivity of a Uruguayan grassland. Plant Ecology 179:83-91. https://doi.org/10.1007/s11258-004-5800-5.

Altesor, A., G. Piñeiro, F. Lezama, R. B. Jackson, M. Sarasola, and J. M. Paruelo. 2006. Ecosystem changes associated with grazing in subhumid South American grasslands. Journal of Vegetation Science 17:323-332. https://doi.org/10.1111/j.1654-1103.2006.tb02452.x.

Altesor, A., F. Gallego, M. Ferrón, F. Pezzani, L. López-Mársico, F. Lezama, S. Baeza, M. Pereira, B. Costa, and J. M. Paruelo. 2019. Inductive Approach To Build State-and-Transition Models for Uruguayan Grasslands. Rangeland Ecology and Management 72:1005-1016. https://doi.org/10.1016/j.rama.2019.06.004.

Andrade, B. O., E. Marchesi, S. Burkart, R. B. Setubal, F. Lezama, S. Perelman, A. A. Schneider, R. Trevisan, G. E. Overbeck, and I. I. Boldrini. 2018. Vascular plant species richness and distribution in the Río de la Plata grasslands. Botanical Journal of the Linnean Society 188:250-256. https://doi.org/10.1093/botlinnean/boy063.

Baeza, S., and J. M. Paruelo. 2018. Spatial and temporal variation of human appropriation of net primary production in the Rio de la Plata grasslands. ISPRS Journal of Photogrammetry and Remote Sensing 145:238-249. https://doi.org/10.1016/j.isprsjprs.2018.07.014.

Baeza, S., and J. M. Paruelo. 2020. Land use/land cover change (2000-2014) in the Río de la Plata grasslands: An analysis based on MODIS NDVI time series. Remote Sensing 12:1-22. https://doi.org/10.3390/rs12030381.

Baldi, G., and J. M. Paruelo. 2008. Land-use and land cover dynamics in South American Temperate grasslands. Ecology and Society 13. http://doi.org/10.5751/ES-02481-130206.

Bates, D., M. Mächler, B. M. Bolker, and S. C. Walker. 2015. Fitting linear mixed-effects models using lme4. Journal of Statistical Software 67. https://doi.org/10.18637/jss.v067.i01.

Behling, H., and V. D. Pillar. 2007. Late Quaternary vegetation, biodiversity and fire dynamics on the southern Brazilian highland and their implication for conservation and management of modern Araucaria forest and grassland ecosystems. Philosophical Transactions of the Royal Society B 362:243-251. https://doi.org/10.1098/rstb.2006.1984.

Behling, H., V. D. Pillar, and S. G. Bauermann. 2005. Late Quaternary grassland (Campos), gallery forest, fire and climate dynamics, studied by pollen, charcoal and multivariate analysis of the São Francisco de Assis core in western Rio Grande do Sul (southern Brazil). Review of Palaeobotany and Palynology 133:235-248. https://doi.org/10.1016/j.revpalbo.2004.10.004.

Behling, H., V. D. Pillar, S. C. Müller, and G. E. Overbeck. 2007. Late‐Holocene fire history in a forest‐grassland mosaic in southern Brasil: Implications for conservation. Applied Vegetation Science 10:81-90. https://doi.org/10.1111/j.1654-109X.2007.tb00506.x.

Bengtsson, J., J. M. Bullock, B. Egoh, C. Everson, T. Everson, T. O’Connor, P. J. O’Farrell, H. G. Smith, and R. Lindborg. 2019. Grasslands—more important for ecosystem services than you might think. Ecosphere 10. https://doi.org/10.1002/ecs2.2582.

Bernardi, R. E., M. Holmgren, M. Arim, and M. Scheffer. 2016. Why are forests so scarce in subtropical South America? The shaping roles of climate, fire and livestock. Forest Ecology and Management 363:212-217. https://doi.org/10.1016/j.foreco.2015.12.032.

Bernardi, R. E., M. Buddeberg, M. Arim, and M. Holmgren. 2019a. Forests expand as livestock pressure declines in subtropical South America. Ecology and Society 24. https://doi.org/10.5751/ES-10688-240219.

Bernardi, R. E., A. Staal, C. Xu, M. Scheffer, and M. Holmgren. 2019b. Livestock Herbivory Shapes Fire Regimes and Vegetation Structure Across the Global Tropics. Ecosystems 22:1457-1465. https://doi.org/10.1007/s10021-019-00349-x.

Bond, W. J. 2005. Large parts of the world are brown or black: A different view on the “Green World” hypothesis. Journal of Vegetation Science 16: 261-266. https://doi.org/10.1111/j.1654-1103.2005.tb02364.x.

Bond, W. J. 2019. Open Ecosystems ecology and evolution beyond the forest edge. Oxford, Oxford, UK. https://doi.org/10.1093/oso/9780198812456.001.0001.

Brazeiro, A., P. Brussa, and C. Toranza. 2018. Efectos del ganado en el ecotono bosque-pastizal en paisajes serranos de Uruguay. Ecosistemas 27:14-23. https://doi.org/10.7818/ECOS.1470.

Carlucci, M. B., L. da S. Duarte, and V. D. Pillar. 2011. Nurse rocks influence forest expansion over native grassland in southern Brazil. Journal of Vegetation Science 22:111-119. https://doi.org/10.1111/j.1654-1103.2010.01229.x.

Carlucci, M. B., A. L. Luza, S. M. Hartz, and L. D. S. Duarte. 2016. Forests, shrublands and grasslands in southern Brazil are neglected and have specific needs for the in conservation. Reply to Overbeck et al. Natureza e Conservação 14:155-157. https://doi.org/10.1016/j.ncon.2016.08.001.

Celaya, R., M. Oliván, L. M. M. Ferreira, A. Martínez, U. García, and K. Osoro. 2007. Comparison of grazing behaviour, dietary overlap and performance in non-lactating domestic ruminants grazing on marginal heathland areas. Livestock Science 106:271-281. https://doi.org/10.1016/j.livsci.2006.08.013.

Clarke, K. R. 1993. Non‐parametric multivariate analyses of changes in community structure. Australian Journal of Ecology 18:117-143. https://doi.org/10.1111/j.1442-9993.1993.tb00438.x.

Cuello, N., L. López-Mársico, and C. Rodríguez. 2020. Field burn versus fire-related cues: germination from the soil seed bank of a South American temperate grassland. Seed Science Research 30:206-214. https://doi.org/10.1017/S0960258520000288.

Dantas, V. de L., M. Hirota, R. S. Oliveira, and J. G. Pausas. 2016. Disturbance maintains alternative biome states. Ecology Letters 19:12-19. https://doi.org/10.1111/ele.12537.

De Cáceres, M., and P. Legendre. 2009 Associations between species and groups of sites: indices and statistical inference. Ecology 90(12): 3566-3574. https://doi.org/10.1890/08-1823.1.

Decreto Nº 60/020. 2020. URL: impo.com.uy/bases/decretos/60-2020.

Dinama-Mvotma. 2019. Proceso de ingreso de Quebrada de los Cuervos y Sierras del Yerbal al Sistema Nacional de Áreas Naturales Protegidas Proyecto de Selección y delimitación del área. URL: tinyurl.com/yc378f3w.

Dufrêne, M., and P. Legendre. 1997. Species assemblages and indicator species: the need for a flexible asymmetrical approach. Ecological Monographs 67:345-366. https://doi.org/10.1890/0012-9615(1997)067[0345:SAAIST]2.0.CO;2.

Durán, A., A. Califra, and J. H. Molfino. 1999. Suelos del Uruguay según Soil Taxonomy.

Eldridge, D. J., M. A. Bowker, F. T. Maestre, E. Roger, J. F. Reynolds, and W. G. Whitford. 2011. Impacts of shrub encroachment on ecosystem structure and functioning: Towards a global synthesis. Ecology Letters 14:709-722. https://doi.org/10.1111/j.1461-0248.2011.01630.x.

Ellis, E. C. 2011. Anthropogenic transformation of the terrestrial biosphere. Philosophical Transactions of the Royal Society A 369:1010-1035. https://doi.org/10.1098/rsta.2010.0331.

Epstein, H. E., R. A. Gill, J. M. Paruelo, W. K. Lauenroth, G. J. Jia, and I. C. Burke. 2002. The relative abundance of three plant functional types in temperate grasslands and shrublands of North and South America: Effects of projected climate change. Journal of Biogeography 29:875-888. https://doi.org/10.1046/j.1365-2699.2002.00701.x.

Etchebarne, V., and A. Brazeiro. 2016. Effects of livestock exclusion in forests of Uruguay: Soil condition and tree regeneration. Forest Ecology and Management 362:120-129. https://doi.org/10.1016/j.foreco.2015.11.042.

Fan, Y., G. Miguez-Macho, E. G. Jobbágy, R. B. Jackson, and C. Otero-Casal. 2017. Hydrologic regulation of plant rooting depth. Proceedings of the National Academy of Sciences of the United States of America 114:10572-10577. https://doi.org/10.1073/pnas.1712381114.

Fisher, N. I. 1983. Graphical Methods in Nonparametric Statistics: A Review and Annotated Bibliography. International Statistical Review 51:25-58. https://doi.org/10.2307/1402730.

Gallego, F. 2013. Servicios ecosistémicos del pastizal: el seguimiento de un área protegida como sistema de referencia. Tesis Maestría en Ciencias Ambientales, Facultad de Ciencias, Universidad de la República.

Gallego, F., J. M. Paruelo, S. Baeza, and A. Altesor. 2020. Distinct ecosystem types respond differentially to grazing exclosure. Austral Ecology 45:548-556. https://doi.org/10.1111/aec.12870.

Gang, C., W. Zhou, Y. Chen, Z. Wang, Z. Sun, J. Li, J. Qi, and I. Odeh. 2014. Quantitative assessment of the contributions of climate change and human activities on global grassland degradation. Environmental Earth Sciences 72:4273-4282. https://doi.org/10.1007/s12665-014-3322-6.

Gautreau, P., and F. Lezama. 2009. Clasificación florística de los bosques y arbustales de las sierras del Uruguay. Ecología Austral 19:81-92.

Gibson, D. J., and J. A. Newman. 2019. Grasslands and Climate Change an overview. Pp. 3-18 en D. J. Gibson and J. A. Newman (eds.). Grasslands and Climate Change. Cambridge University Press, UK. https://doi.org/10.1017/9781108163941.003.

Graesser, J., T. M. Aide, H. R. Grau, and N. Ramankutty. 2015. Cropland/pastureland dynamics and the slowdown of deforestation in Latin America. Environmental Research Letters 10. https://doi.org/10.1088/1748-9326/10/3/034017.

Guido, A., E. Salengue, and A. Dresseno. 2017. Effect of shrub encroachment on vegetation communities in Brazilian forest-grassland mosaics. Perspectives in Ecology and Conservation 15:52-55. https://doi.org/10.1016/j.pecon.2016.11.002.

Haretche, F., P. Mai, and A. Brazeiro. 2012. Flora lenhosa do Uruguai: Inventário e implicação na região Pampeana. Acta Botanica Brasilica 26:537-552. https://doi.org/10.1590/S0102-33062012000300004.

Hayes, G. F., and K. D. Holl. 2003. Cattle Grazing Impacts on Annual Forbs and Vegetation Composition of Mesic Grasslands in California. Conservation Biology 17:1694-1702. https://doi.org/10.1111/j.1523-1739.2003.00281.x.

Hirota, M., M. Holmgren, E. H. van Nes, and M. Scheffer. 2011. Global Resilience of Tropical Forest and Savanna to Critical Transitions. Science 334:232-235. https://doi.org/10.1126/science.1210657.

Hofmann, R. R. 1989. Evolutionary steps of ecophysiological adaptation and diversification of ruminants: a comparative view of their digestive system. Oecologia 78:443-457. https://doi.org/10.1007/BF00378733.

Holdridge, L. R. 1947. Determination of World Plant Formations From Simple Climatic Data. Science 105:367-368. https://doi.org/10.1126/science.105.2727.367.

Holmgren, M., C. Y. Lin, J. E. Murillo, A. Nieuwenhuis, J. Penninkhof, N. Sanders, T. van Bart, H. van Veen, H. Vasander, M. E. Vollebregt, and J. Limpens. 2015. Positive shrub-tree interactions facilitate woody encroachment in boreal peatlands. Journal of Ecology 103:58-66. https://doi.org/10.1111/1365-2745.12331.

IPBES. 2018. The IPBES regional assessment report on biodiversity and ecosystem services for the Americas. Rice, J., C. S. Seixas, M. E. Zaccagnini, M. Bedoya-Gaitán, and N. Valderrama (eds.). Secretariat of the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services, Bonn, DK.

InuMet. 2020. Tablas estadísticas. URL: tinyurl.com/2p8tt7ze.

IUCN. 2000. Guidelines for the Prevention of biodiversity loss caused by alien invasive species.

Kark, S. 2013. Encyclopedia of Sustainability Science and Technology. Pp. 147-160 en R. Leemans (ed.). Encyclopedia of Sustainability Science and Technology. Springer, New York, USA.

Kuznetsova, A., P. B. Brockhoff, and R. H. B. Christensen. 2017. lmerTest Package: Tests in Linear MixedEffects Models. Journal of Statistical Software 82(13):1-26. https://doi.org/10.18637/jss.v082.i13.

Lapetina, J. 2012. Guía de buenas prácticas ganaderas para el manejo y conservación de pastizales naturales en áreas protegidas. Sistema Nacional de Áreas Protegidas - Ministerio de Vivienda, Ordenamiento territorial y Medio Ambiente. URL: tinyurl.com/4v4k4sc3.

Legendre, P., and L. Legendre. 2012. Numerical Ecology. Third Engl. Elsevier, Oxford, UK.

Leite, M. B., R. O. Xavier, P. T. S. Oliveira, F. K. G. Silva, and D. M. Silva Matos. 2018. Groundwater depth as a constraint on the woody cover in a Neotropical Savanna. Plant and Soil 426. https://doi.org/10.1007/s11104-018-3599-4.

Lesica, P. 2015. Monitoring Plants at Ecotones for Effects of Environmental Change. Natural Areas Journal 35:485-487. https://doi.org/10.3375/043.035.0315.

Lezama, F., S. Baeza, A. Altesor, A. Cesa, E. J. Chaneton, and J. M. Paruelo. 2014. Variation of grazing-induced vegetation changes across a large-scale productivity gradient. Journal of Vegetation Science 25:8-21. https://doi.org/10.1111/jvs.12053.

Lezama, F., and J. M. Paruelo. 2016. Disentangling grazing effects: trampling, defoliation and urine deposition. Applied Vegetation Science 19:557-566. https://doi.org/10.1111/avsc.12250.

Lezama, F., M. Pereira, A. Altesor, and J. M. Paruelo. 2019. Grasslands of Uruguay: Classification based on vegetation plots. Phytocoenologia 49:211-229. https://doi.org/10.1127/phyto/2019/0215.

Lloyd, K. M., A. A. M. McQueen, B. J. Lee, R. C. B. Wilson, S. Walker, and J. B. Wilson. 2000. Evidence on ecotone concepts from switch, environmental and anthropogenic ecotones. Journal of Vegetation Science 11:903-910. https://doi.org/10.2307/3236560.

López-Márisco, L., F. Lezama, and A. Altesor. 2020. Heterogeneity decreases as time since fire increases in a South American grassland. Applied Vegetation Science 24:e12521. https://doi.org/10.1111/avsc.12521.

López-Mársico, L., L. Farías-Moreira, F. Lezama, A. Altesor, and C. Rodríguez. 2019. Light intensity triggers different germination responses to fire-related cues in temperate grassland species. Folia Geobotanica 54:53-63. https://doi.org/10.1007/s12224-019-09336-5.

Luza, A. L., M. B. Carlucci, S. M. Hartz, and L. D. S. Duarte. 2014. Moving from forest vs. grassland perspectives to an integrated view towards the conservation of forest-grassland mosaics. Natureza and Conservação 12:166-169. https://doi.org/10.1016/j.ncon.2014.09.005.

van der Maarel, E. 2007. Transformation of cover-abundance values for appropriate numerical treatment - Alternatives to the proposals by Podani. Journal of Vegetation Science 18:767. https://doi.org/10.1111/j.1654-1103.2007.tb02592.x.

Mantyka-Pringle, C. S., T. G. Martin, and J. R. Rhodes. 2012. Interactions between climate and habitat loss effects on biodiversity: A systematic review and meta-analysis. Global Change Biology 18:1239-1252. https://doi.org/10.1111/j.1365-2486.2011.02593.x.

Martínez, M., and M. Pereira. 2011. Pautas para el manejo del campo natural. Instituto Plan Agropecuario, Uruguay. URL: tinyurl.com/2r67nysb.

Mesa de Ganadería sobre Campo Natural. 2016. Curso Producción animal sostenible en pastoreo sobre campo natural. Uy. URL: tinyurl.com/bdsnxy76.

Modernel, P., W. A. H. Rossing, M. Corbeels, S. Dogliotti, V. Picasso, and P. Tittonell. 2016. Land use change and ecosystem service provision in Pampas and Campos grasslands of southern South America. Environmental Research Letters 11:113002. https://doi.org/10.1088/1748-9326/11/11/113002.

Moncrieff, G. R., W. J. Bond, and S. I. Higgins. 2016. Revising the biome concept for understanding and predicting global change impacts. Journal of Biogeography 43:863-873. https://doi.org/10.1111/jbi.12701.

Müller, S. C., G. E. Overbeck, J. Pfadenhauer, and V. D. Pillar. 2007. Plant functional types of woody species related to fire disturbance in forest-grassland ecotones. Plant Ecology 189:1-14. https://doi.org/10.1007/s11258-006-9162-z.

Müller, S. C., G. E. Overbeck, J. Pfadenhauer, and V. D. Pillar. 2012. Woody species patterns at forest - grassland boundaries in southern Brazil. Flora 207:586-598. https://doi.org/10.1016/j.flora.2012.06.012.

Murphy, B. P., A. N. Andersen, and C. L. Parr. 2016. The underestimated biodiversity of tropical grassy biomes. Philosophical Transactions of the Royal Society B: Biological Sciences 371. https://doi.org/10.1098/rstb.2015.0319.

Murphy, B. P., and D. M. J. S. Bowman. 2012. What controls the distribution of tropical forest and savanna? Ecology Letters 15:748-758. https://doi.org/10.1111/j.1461-0248.2012.01771.x.

Newbold, T. 2018. Future effects of climate and land-use change on terrestrial vertebrate community diversity under different scenarios. Proceedings of the Royal Society B: Biological Sciences 285. https://doi.org/10.1098/rspb.2018.0792.

Newton, A. C. 2007. Forest Ecology and Conservation. First edition. Oxford University Press, New York, USA.

Oksanen, J., F. Guillaume Blanchet, M. Friendly, R. Kindt, P. Legendre, D. McGlinn, P. R. Minchin, R. B. O’Hara, G. L. Simpson, P. Solymos, M. H. H. Stevens, E. Szoecs, and H. Wagner 2019. vegan: Community Ecology Package. R package version 2.5-5. URL: CRAN.R-project.org/package=vegan.

Oliveira-Filho, A. T., J. C. Budke, J. A. Jarenkow, P. V. Eisenlohr, and D. R. M. Neves. 2015. Delving into the variations in tree species composition and richness across South American subtropical Atlantic and Pampean forests. Journal of Plant Ecology 8:242-260. https://doi.org/10.1093/jpe/rtt058.

Oliveira, J. M., and V. D. Pillar. 2004. Vegetation dynamics on mosaics of Campos and Araucaria forest between 1974 and 1999 in Southern Brazil. Community Ecology 5:197-202. https://doi.org/10.1556/ComEc.5.2004.2.8.

O’Mara, F. P. 2012. The role of grasslands in food security and climate change. Annals of Botany 110:1263-1270. https://doi.org/10.1093/aob/mcs209.

Overbeck, G. E., P. M. A. Ferreira, and V. D. Pillar. 2016. Conservation of mosaics calls for a perspective that considers all types of mosaic-patches. Reply to Luza et al. Natureza e Conservação 14:152-154. https://doi.org/10.1016/j.ncon.2016.05.002.

Overbeck, G. E., S. C. Müller, A. Fidelis, J. Pfadenhauer, V. D. Pillar, C. C. Blanco, I. I. Boldrini, R. Both, and E. D. Forneck. 2007. Brazil’s neglected biome: The South Brazilian Campos. Perspectives in Plant Ecology, Evolution and Systematics 9:101-116. https://doi.org/10.1016/j.ppees.2007.07.005.

Overbeck, G. E., and J. Pfadenhauer. 2007. Adaptive strategies in burned subtropical grassland in southern Brazil. Flora: Morphology, Distribution, Functional Ecology of Plants 202:27-49. https://doi.org/10.1016/j.flora.2005.11.004.

Oyarzabal, M., B. Andrade, V. D. Pillar, and J. Paruelo. 2020. Temperate Subhumid Grasslands of Southern South America. En M. Goldstein and D. Della Sala (eds.). Encyclopedia of the World’s Biomes. Elsevier Inc. https://doi.org/10.1016/B978-0-12-409548-9.12132-3.

Panario, D., O. Gutiérrez, L. S. Bettucci, E. Peel, P. Oyhantçabal, and J. Rabassa. 2015. Ancient landscapes of Uruguay. Pp. 161-199 en J. Rabassa and C. Ollier (eds.). Gondwana Landscapes in southern South America. Springer, New York, USA. https://doi.org/10.1007/978-94-007-7702-6_8.

Paradis E., and K. Schliep. 2019. ape 5.0: an environment for modern phylogenetics and evolutionary analyses in R. Bioinformatics 35: 526-528 https://doi.org/10.1093/bioinformatics/bty633.

Paruelo, J. M., E. G. Jobbágy, M. Oesterheld, R. A. Golluscio, and M. R. Aguiar. 2007. The Grasslands and Steppes of Patagonia and the Río de la Plata Plains. Pp. 232-249 en T. Veblen, K. Young and A. Orme (eds.). The Physical Geography of South America. Oxford University Press. https://doi.org/10.1093/oso/9780195313413.003.0022.

Pausas, J. G., and W. J. Bond. 2020. Alternative Biome States in Terrestrial Ecosystems. Trends in Plant Science 25:250-263. https://doi.org/10.1016/j.tplants.2019.11.003.

Peters, D. P. C., J. R. Gosz, W. T. Pockman, E. E. Small, R. R. Parmenter, S. L. Collins, and E. Muldavin. 2006. Integrating patch and boundary dynamics to understand and predict biotic transitions at multiple scales. Landscape Ecology 21:19-33. https://doi.org/10.1007/s10980-005-1063-3.

Pillar De Patta, V., and F. de Quadros. 1997. Grassland-forest boundaries in Southern Brazil. COENOSES 12:119-126.

Prache, S., I. J. Gordon, and A. J. Rook. 1998. Foraging behaviour and diet selection in domestic herbivores. Animal Research 47:335-345. https://doi.org/10.1051/animres:19980502.

R Core Team. 2019. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL: R-project.org.

Ratajczak, Z., J. B. Nippert, J. M. Briggs, and J. M. Blair. 2014. Fire dynamics distinguish grasslands , shrublands and woodlands as alternative attractors in the Central Great Plains of North America 102:1374-1385. https://doi.org/10.1111/1365-2745.12311.

Rodríguez, C., E. Leoni, F. Lezama, and A. Altesor. 2003. Temporal trends in species composition and plant traits in natural grasslands of Uruguay. Journal of Vegetation Science 14:433-440. https://doi.org/10.1111/j.1654-1103.2003.tb02169.x.

Rolhauser, A. G., and W. B. Batista. 2014. From pattern to process: estimating expansion rates of a forest tree species in a protected palm savanna. Landscape Ecology 29:919-931. https://doi.org/10.1007/s10980-014-0029-8.

RStudio Team. 2016. RStudio: Integrated Development for R. RStudio, Inc., Boston, MA. URL: rstudio.com.

Sankey, T. T. 2012. Woody-Herbaceous-livestock Species Interactions. Pp. 89-114 en R. W. Myster (ed.). Ecotones Between Forest and Grassland. Springer Science+Business Media, New York, USA. https://doi.org/10.1007/978-1-4614-3797-0_4.

Scheffer, M., S. Carpenter, J. A. Foley, C. Folke, and B. Walker. 2001. Catastrophic shifts in ecosystems. Nature 413:591-596. https://doi.org/10.1038/35098000.

Schinestsck, C. F., S. C. Müller, and V. D. Pillar. 2019. Woody species patterns linked to the process of araucaria forest expansion over native grasslands excluded from management. Neotropical Biology and Conservation 14:411-429. https://doi.org/10.3897/neotropical.14.e47885.

Seddon, A. W. R., M. Macias-Fauria, P. R. Long, D. Benz, and K. J. Willis. 2016. Sensitivity of global terrestrial ecosystems to climate variability. Nature 531: 229-232. https://doi.org/10.1038/nature16986.

Silveira, F. A. O., A. J. Arruda, W. Bond, G. Durigan, A. Fidelis, K. Kirkman, R. S. Oliveira, G. E. Overbeck, J. B. B. Sansevero, F. Siebert, S. J. Siebert, T. P. Young, and E. Buisson. 2020. Myth-busting tropical grassy biome restoration. Restoration Ecology 28:1067-1073. https://doi.org/10.1111/rec.13202.

Silveira, F. A. O., C. A. Ordóñez‐Parra, L. C. Moura, I. B. Schmidt, A. N. Andersen, W. Bond, E. Buisson, G. Durigan, A. Fidelis, R. S. Oliveira, C. Parr, L. Rowland, J. W. Veldman, and R. T. Pennington. 2021. Biome Awareness Disparity is BAD for tropical ecosystem conservation and restoration. Journal of Applied Ecology 00:1-9. https://doi.org/10.1111/1365-2664.14060.

Song, X. P., M. C. Hansen, S. V. Stehman, P. V. Potapov, A. Tyukavina, E. F. Vermote, and J. R. Townshend. 2018. Global land change from 1982 to 2016. Nature 560:639-643. https://doi.org/10.1038/s41586-018-0411-9.

Soriano, A. 1991. Rio de la Plata Grasslands. Pp. 367-407 en R. T. Coupland (ed.). Natural Grasslands: introduction and Western Hemisphere. Elsevier, Amsterdam, NL.

Soutullo, A., C. Clavijo, and J. A. Martínes-Lafranco (eds.). 2013. Especies prioritarias para la conservación en Uruguay. snap/dinama/mvotma y dicyt/mec, Montevideo, Uy.

Staver, A. C., S. Archibald, and S. A. Levin. 2011. The global extent and determinants of savanna and forest as alternative biome states. Science 334:230-232. https://doi.org/10.1126/science.1210465.

The World Bank Environment Department. 2010. Convenient Solutions to an Inconvenient Truth : Approaches to Climate Change. Page World. The World Bank, Washington, USA.

Tiscornia, G., M. Jaurena, and W. Baethgen. 2019. Drivers, process, and consequences of native grassland degradation: Insights from a literature review and a survey in Río de la Plata grasslands. Agronomy 9:8-12. https://doi.org/10.3390/agronomy9050239.

United States Department of Agriculture. 1999. Soil Taxonomy A Basic System of Soil Classification for Making and Interpreting Soil Surveys. Second edition. Washington, USA.

Veldman, J. W., E. Buisson, G. Durigan, G. W. Fernandes, S. Le Stradic, G. Mahy, D. Negreiros, G. E. Overbeck, R. G. Veldman, N. P. Zaloumis, F. E. Putz, and W. J. Bond. 2015. Toward an old-growth concept for grasslands, savannas, and woodlands. Frontiers in Ecology and the Environment 13:154-162. https://doi.org/10.1890/140270.

Venables, W. N., and B. D Ripley. 2002. Modern Applied Statistics with S. Springer, New York, USA. https://doi.org/10.1007/978-0-387-21706-2.

Volante, J., J. Mosciaro, M. Morales Poclava, L. Vale, S. Castrillo, J. Sawchik, G. Tiscornia, M. Fuente, I. Maldonado, A. Vega, R. Trujillo, L. Cort’z, and J. Paruelo. 2015. Expansión agrícola en Argentina, Bolivia, Paraguay, Uruguay y Chile entre 2000-2010. Caracterización espacial mediante series temporales de índices de vegetación. Revista de Investigaciones Agropecuarias 41:179-191.

Watt, A. S. 1947. Pattern and process on the Plant community. The Journal of Ecology 35:1-22. https://doi.org/10.2307/2256497.

Whittaker, R. H. 1975. Communities and ecosystems. Page Environmental Biology. Second edition. MacMillan Publishing Co., INC, New York, US.

Zuur, A. F., E. N. Ieno, N. Walker, A. A. Saveliev, and G. M. Smith. 2009. Mixed effects models and extensions in ecology with R. Springer, New York, USA. https://doi.org/10.1007/978-0-387-87458-6.