{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,1]],"date-time":"2026-06-01T15:28:05Z","timestamp":1780327685535,"version":"3.54.1"},"reference-count":86,"publisher":"MDPI AG","issue":"8","license":[{"start":{"date-parts":[[2021,4,14]],"date-time":"2021-04-14T00:00:00Z","timestamp":1618358400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","award":["SE2383\/5-1"],"award-info":[{"award-number":["SE2383\/5-1"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100002850","name":"Fondo Nacional de Desarrollo Cient\u00edfico y Tecnol\u00f3gico","doi-asserted-by":"publisher","award":["11181140"],"award-info":[{"award-number":["11181140"]}],"id":[{"id":"10.13039\/501100002850","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Remote Sensing"],"abstract":"<jats:p>The structural complexity of the understory layer of forests or shrub layer vegetation in open shrublands affects many ecosystem functions and services provided by these ecosystems. We investigated how the basal area of the overstory layer, annual and seasonal precipitation, annual mean temperature, as well as light availability affect the structural complexity of the understory layer along a gradient from closed forests to open shrubland with only scattered trees. Using terrestrial laser scanning data and the understory complexity index (UCI), we measured the structural complexity of sites across a wide range of precipitation and temperature, also covering a gradient in light availability and basal area. We found significant relationships between the UCI and tree basal area as well as canopy openness. Structural equation models (SEMs) confirmed significant direct effects of seasonal precipitation on the UCI without mediation through basal area or canopy openness. However, annual precipitation and temperature effects on the UCI are mediated through canopy openness and basal area, respectively. Understory complexity is, despite clear dependencies on the available light and overall stand density, significantly and directly driven by climatic parameters, particularly the amount of precipitation during the driest month.<\/jats:p>","DOI":"10.3390\/rs13081513","type":"journal-article","created":{"date-parts":[[2021,4,14]],"date-time":"2021-04-14T15:30:39Z","timestamp":1618414239000},"page":"1513","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":17,"title":["Quantifying Understory Complexity in Unmanaged Forests Using TLS and Identifying Some of Its Major Drivers"],"prefix":"10.3390","volume":"13","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-4131-9424","authenticated-orcid":false,"given":"Dominik","family":"Seidel","sequence":"first","affiliation":[{"name":"Silviculture and Forest Ecology of the Temperate Zones, Faculty of Forest Sciences, University of G\u00f6ttingen, B\u00fcsgenweg 1, 37077 G\u00f6ttingen, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peter","family":"Annigh\u00f6fer","sequence":"additional","affiliation":[{"name":"Forest and Agroforest Systems, Technical University of Munich, Hans-Carl-v.-Carlowitz-Platz 2, 85354 Freising, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4235-0135","authenticated-orcid":false,"given":"Christian","family":"Ammer","sequence":"additional","affiliation":[{"name":"Silviculture and Forest Ecology of the Temperate Zones, Faculty of Forest Sciences, University of G\u00f6ttingen, B\u00fcsgenweg 1, 37077 G\u00f6ttingen, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Martin","family":"Ehbrecht","sequence":"additional","affiliation":[{"name":"Silviculture and Forest Ecology of the Temperate Zones, Faculty of Forest Sciences, University of G\u00f6ttingen, B\u00fcsgenweg 1, 37077 G\u00f6ttingen, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0129-3381","authenticated-orcid":false,"given":"Katharina","family":"Willim","sequence":"additional","affiliation":[{"name":"Silviculture and Forest Ecology of the Temperate Zones, Faculty of Forest Sciences, University of G\u00f6ttingen, B\u00fcsgenweg 1, 37077 G\u00f6ttingen, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jan","family":"Bannister","sequence":"additional","affiliation":[{"name":"Instituto Forestal, INFOR, Oficina Chilo\u00e9, Castro 7770223, Chile"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7119-8558","authenticated-orcid":false,"given":"Daniel P.","family":"Soto","sequence":"additional","affiliation":[{"name":"Departamento de Ciencias Naturales y Tecnolog\u00eda, Universidad de Ays\u00e9n, Coyhaique 5950000, Chile"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2021,4,14]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"823","DOI":"10.1051\/forest:2003077","article-title":"Effects of tree species on understory vegetation and environmental conditions in temperate forests","volume":"60","author":"Augusto","year":"2003","journal-title":"Ann. For. Sci."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1016\/S0378-1127(01)00549-7","article-title":"Rationale and methods for conserving biodiversity in plantation forests","volume":"155","author":"Hartley","year":"2002","journal-title":"For. Ecol. Manag."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"864","DOI":"10.1890\/1051-0761(1999)009[0864:PDIMFU]2.0.CO;2","article-title":"Plant diversity in managed forests: Understory responses to thinning and fertilization","volume":"9","author":"Thomas","year":"1999","journal-title":"Ecol. Appl."},{"key":"ref_4","unstructured":"Kimmins, J.P. (2004). The Biogeochemical Cycle: Nutrient Cycling Within Ecosystems. Forest Ecology: A Foundation for Sustainable Forest Management and Environmental Ethics in Forestry, Prentice Hall. [3rd ed.]."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1615","DOI":"10.1111\/1365-2664.12902","article-title":"Effects of deer on woodland structure revealed through terrestrial laser scanning","volume":"54","author":"Eichhorn","year":"2017","journal-title":"J. Appl. Ecol."},{"key":"ref_6","first-page":"262","article-title":"Understory Plants in Temperate Forests","volume":"1","author":"Antos","year":"2009","journal-title":"For. For. Plants"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"241","DOI":"10.4067\/S0717-92002006000300003","article-title":"Patrones de regeneraci\u00f3n en renovales de Drimys winteri en el centro-norte de la Isla de Chilo\u00e9: Cambios de acuerdo al tama\u00f1o y la densidad relativa","volume":"27","author":"Soto","year":"2006","journal-title":"Bosque"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1016\/0378-1127(93)90144-C","article-title":"Long-and short-term effects of alternative cutting regimes on economic returns and ecological diversity in mixed-species forests","volume":"58","author":"Lu","year":"1993","journal-title":"For. Ecol. Manag."},{"key":"ref_9","first-page":"51","article-title":"Densidad de pl\u00e1ntulas de acuerdo a la estructura, dominancia y cobertura del sotobosque en bosques siempreverdes adultos en la cordillera de la Costa de Chile","volume":"78","author":"Donoso","year":"2005","journal-title":"Rev. Chil. Hist. Nat."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"26","DOI":"10.1016\/j.foreco.2019.05.004","article-title":"Regeneration niches in Nothofagus-dominated old-growth forests after partial disturbance: Insights to overcome arrested succession","volume":"445","author":"Soto","year":"2019","journal-title":"For. Ecol. Manag."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1126\/science.aai8291","article-title":"Plant-soil feedback and the maintenance of diversity in Mediterranean-climate shrublands","volume":"355","author":"Teste","year":"2017","journal-title":"Science"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"184","DOI":"10.1007\/s10310-011-0257-6","article-title":"The ecology of tree regeneration in mature and old forests: Combined knowledge for sustainable forest management","volume":"16","author":"Liira","year":"2011","journal-title":"J. For. Res."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1016\/j.foreco.2018.08.039","article-title":"A quantitative comparison of the structural complexity of managed, lately unmanaged and primary European beech (Fagus sylvatica L.) forests","volume":"430","author":"Stiers","year":"2018","journal-title":"For. Ecol. Manag."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s10342-005-0089-z","article-title":"Trends in North American and European regeneration research under the ecosystem management paradigm","volume":"126","author":"Puettmann","year":"2007","journal-title":"Eur. J. For. Res."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"967","DOI":"10.1111\/1365-2664.12989","article-title":"Topsoil removal through scarification improves natural regeneration in high-graded Nothofagus old-growth forests","volume":"55","author":"Soto","year":"2018","journal-title":"J. Appl. Ecol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"23","DOI":"10.4067\/S0717-92002013000100004","article-title":"Early performance of planted Nothofagus dombeyi and Nothofagus alpina in response to light availability and gap size in a high-graded forest in the south-central Andes of Chile","volume":"34","author":"Donoso","year":"2013","journal-title":"Bosque"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1","DOI":"10.2307\/2963479","article-title":"Forest models defined by field measurements: Estimation, error analysis and dynamics","volume":"66","author":"Pacala","year":"1996","journal-title":"Ecol. Monogr."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1016\/j.foreco.2016.11.016","article-title":"Light and nitrogen interact to influence regeneration in old-growth Nothofagus-dominated forests in south-central Chile","volume":"384","author":"Soto","year":"2017","journal-title":"For. Ecol. Manag."},{"key":"ref_19","first-page":"1611","article-title":"Nitrogen-induced changes in seedling regeneration and dynamics of mixed coniferbroad-leaved forests","volume":"12","author":"Catovsky","year":"2002","journal-title":"Ecol. Appl."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"690","DOI":"10.3389\/fpls.2019.00690","article-title":"Advanced aboveground spatial analysis as proxy for the competitive environment affecting sapling development","volume":"10","author":"Seidel","year":"2019","journal-title":"Front. Plant Sci."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1016\/0378-1127(95)97453-Y","article-title":"Effects of soil water content, fertilization, light, weed competition and seedbed type on natural regeneration of beech (Fagus sylvatica)","volume":"72","author":"Madsen","year":"1995","journal-title":"For. Ecol. Manag."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"387","DOI":"10.1016\/S0378-1127(03)00085-9","article-title":"Effects of nitrogen and phosphorus fertilization on the survival and recruitment of seedlings of dominant tree species in two abandoned tropical dry forests in Yucat\u00e1n, Mexico","volume":"182","author":"Ceccon","year":"2003","journal-title":"For. Ecol. Manag."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"485","DOI":"10.1046\/j.1466-822X.2002.00306.x","article-title":"Climate change-integrated conservation strategies","volume":"11","author":"Hannah","year":"2002","journal-title":"Glob. Ecol. Biogeogr."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"503","DOI":"10.1111\/j.1744-7429.2009.00613.x","article-title":"Species-specific growth responses to climate variations in understory trees of a Central African rain forest","volume":"42","author":"Couralet","year":"2010","journal-title":"Biotropica"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1543","DOI":"10.1111\/j.1365-2486.2008.01584.x","article-title":"Mixed-conifer understory response to climate change, nitrogen, and fire","volume":"14","author":"Hurteau","year":"2008","journal-title":"Glob. Chang. Biol."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"689","DOI":"10.1007\/s13595-011-0086-y","article-title":"Climatic controls of a keystone understory species, Sasamorpha borealis, and an impact assessment of climate change in Japan","volume":"68","author":"Tsuyama","year":"2011","journal-title":"Ann. For. Sci."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.foreco.2005.08.034","article-title":"Forest and woodland stand structural complexity: Its definition and measurement","volume":"218","author":"McElhinny","year":"2005","journal-title":"For. Ecol. Manag."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"681","DOI":"10.1016\/j.tree.2014.10.005","article-title":"Advances in animal ecology from 3D-LiDAR ecosystem mapping","volume":"29","author":"Davies","year":"2014","journal-title":"Trends Ecol. Evol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1007\/s40725-016-0031-2","article-title":"A review of processes behind diversity\u2014productivity relationships in forests","volume":"2","author":"Forrester","year":"2016","journal-title":"Curr. For. Rep."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"e02864","DOI":"10.1002\/ecy.2864","article-title":"High rates of primary production in structurally complex forests","volume":"100","author":"Gough","year":"2019","journal-title":"Ecology"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1038\/nature07276","article-title":"Old-growth forests as global carbon sinks","volume":"455","author":"Luyssaert","year":"2008","journal-title":"Nature"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1016\/j.biombioe.2012.10.009","article-title":"The potential of terrestrial laser scanning for the estimation of understory biomass in coppice-with-standard systems","volume":"47","author":"Seidel","year":"2012","journal-title":"Biomass Bioenergy"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"196","DOI":"10.1016\/j.foreco.2016.09.036","article-title":"Assessing different components of three-dimensional forest structure with single-scan terrestrial laser scanning: A case study","volume":"381","author":"Seidel","year":"2016","journal-title":"For. Ecol. Manag."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.agrformet.2017.04.012","article-title":"Quantifying stand structural complexity and its relationship with forest management, tree species diversity and microclimate","volume":"242","author":"Ehbrecht","year":"2017","journal-title":"Agric. For. Meteorol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"128","DOI":"10.1002\/ece3.3661","article-title":"A holistic approach to determine tree structural complexity based on laser scanning data and fractal analysis","volume":"8","author":"Seidel","year":"2018","journal-title":"Ecol. Evol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"911","DOI":"10.1007\/s00468-019-01827-4","article-title":"Identifying architectural characteristics that determine tree structural complexity","volume":"33","author":"Seidel","year":"2019","journal-title":"Trees"},{"key":"ref_37","first-page":"75","article-title":"Effect of tree species mixing on stand structural complexity","volume":"93","author":"Juchheim","year":"2020","journal-title":"For. Int. J. For. Res."},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Frey, J., Asbeck, T., and Bauhus, J. (2020). Predicting tree-related microhabitats by multisensor close-range remote sensing structural parameters for the selection of retention elements. Remote Sens., 12.","DOI":"10.3390\/rs12050867"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"107699","DOI":"10.1016\/j.agrformet.2019.107699","article-title":"From tree to stand-level structural complexity\u2014Which properties make a forest stand complex?","volume":"278","author":"Seidel","year":"2019","journal-title":"Agricultural and Forest Meteorology"},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Frey, J., Joa, B., Schraml, U., and Koch, B. (2019). Same viewpoint different perspectives\u2014A comparison of expert ratings with a TLS derived forest stand structural complexity index. Remote Sens., 11.","DOI":"10.3390\/rs11091137"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1038\/s41467-020-20767-z","article-title":"Global patterns and climatic controls of forest structural complexity","volume":"12","author":"Ehbrecht","year":"2021","journal-title":"Nat. Commun."},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Willim, K., Stiers, M., Annigh\u00f6fer, P., Ammer, C., Ehbrecht, M., Kabal, M., Stillhard, J., and Seidel, D. (2019). Assessing understory complexity in beech-dominated forests (Fagus sylvatica L.) in Central Europe\u2014From managed to primary forests. Sensors, 19.","DOI":"10.3390\/s19071684"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"487","DOI":"10.4067\/S0718-58392016000400013","article-title":"Long term climatic trends in Chile and effects on soil moisture and temperature regimes","volume":"76","author":"Stolpe","year":"2016","journal-title":"Chil. J. Agric. Res."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1007\/BF01821439","article-title":"Die Winkelz\u00e4hlprobe","volume":"71","author":"Bitterlich","year":"1952","journal-title":"Forstwissenschaftliches Centralblatt"},{"key":"ref_45","unstructured":"Kramer, H., and Akca, A. (2002). Leitfaden zur Waldmesslehre, JD Sauerl\u00e4nder\u2018s Verlag. 266 S."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"1087","DOI":"10.3390\/f4041087","article-title":"Forest typification to characterize the structure and composition of old-growth evergreen forests on Chiloe Island, North Patagonia (Chile)","volume":"4","author":"Bannister","year":"2013","journal-title":"Forests"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1330","DOI":"10.1139\/cjfr-2016-0167","article-title":"Does site quality affect the additive basal area phenomenon? Results from Chilean old-growth temperate rainforests","volume":"46","author":"Donoso","year":"2016","journal-title":"Can. J. For. Res."},{"key":"ref_48","unstructured":"Donoso, C. (2015). Estructura y Din\u00e1mica de los Bosques del Cono sur de Am\u00e9rica, Edici\u00f3n Universidad Mayor."},{"key":"ref_49","first-page":"675","article-title":"Differential effects of emergent Nothofagus dombeyi on growth and basal area of canopy species in an old-growth temperate rainforest","volume":"18","author":"Donoso","year":"2007","journal-title":"J. Veg. Sci."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"1","DOI":"10.2307\/2259240","article-title":"Structure and dynamics of old-growth Nothofagus forests in the Valdivian Andes, Chile","volume":"68","author":"Veblen","year":"1980","journal-title":"J. Ecol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"1176","DOI":"10.1111\/j.1654-1103.2009.01117.x","article-title":"Replacement patterns and species coexistence in an Andean Araucaria\u2013Nothofagus forest","volume":"20","author":"Fajardo","year":"2009","journal-title":"J. Veg. Sci."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"54","DOI":"10.4067\/S0717-66432016000100007","article-title":"Regeneraci\u00f3n natural y patrones de distribuci\u00f3n espacial de la palma chilena Jubaea chilensis (Molina) Baillon en los bosques mediterr\u00e1neos de Chile central. Gayana","volume":"73","author":"Miranda","year":"2016","journal-title":"Bot\u00e1nica"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"641","DOI":"10.2307\/3236571","article-title":"Variation in vegetation and seed bank in a Chilean semi-arid community affected by ENSO 1997","volume":"11","author":"Arancio","year":"2000","journal-title":"J. Veg. Sci."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"4302","DOI":"10.1002\/joc.5086","article-title":"Worldclim 2: New 1-km spatial resolution climate surfaces for global land areas","volume":"37","author":"Fick","year":"2017","journal-title":"Int. J. Climatol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"623","DOI":"10.1002\/hyp.7850","article-title":"Seasonal and annual throughfall and stemflow in Andean temperate rainforests","volume":"25","author":"Godoy","year":"2011","journal-title":"Hydrol. Process."},{"key":"ref_56","unstructured":"Luzio Leighton, W. (2010). Suelos de Chile, Impresos Maval."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"1187","DOI":"10.1111\/j.1365-2699.2005.01262.x","article-title":"Fire history of Araucaria\u2013Nothofagus forests in Villarrica National Park, Chile","volume":"32","author":"Veblen","year":"2005","journal-title":"J. Biogeogr."},{"key":"ref_58","unstructured":"El\u00f3rtegui, S., and Moreira Mu\u00f1oz, A. (2009). Climatolog\u00eda. Parque Nacional La Campana: Origen de Una Reserva de la Biosfera en Chile Central, Taller La Era. [2nd ed.]."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"500","DOI":"10.1111\/j.1442-9993.2011.02312.x","article-title":"Latitudinal patterns and regionalization of plant diversity along a 4270-km gradient in continental Chile","volume":"37","author":"Bannister","year":"2012","journal-title":"Austral Ecol."},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"McGarigal, K., and Marks, B. (1994). Fragstats: Spatial Pattern Analysis Program for Quantifying Landscape Structure. Vers. 2, U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station.","DOI":"10.2737\/PNW-GTR-351"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"777","DOI":"10.1109\/TGRS.2012.2205003","article-title":"Retrieval of effective leaf area index in heterogeneous forests with terrestrial laser scanning","volume":"51","author":"Zheng","year":"2013","journal-title":"IEEE Transit. Geosci. Remote Sens."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"860","DOI":"10.1016\/j.foreco.2018.10.008","article-title":"Effects of structural heterogeneity on the diurnal temperature range in temperate forest ecosystems","volume":"432","author":"Ehbrecht","year":"2019","journal-title":"For. Ecol. Manag."},{"key":"ref_63","unstructured":"Wood, S.N. (2017). Generalized Additive Models: An Introduction with R, CRC Press. [2nd ed.]. Chapman and Hall\/CRC Texts in Statistical Science."},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Otto, S.A., Diekmann, R., Flinkman, J., Kornilovs, G., and M\u00f6llmann, C. (2014). Habitat heterogeneity determines climate impact on zooplankton community structure and dynamics. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0090875"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"3418","DOI":"10.1890\/03-0755","article-title":"Nonadditive effects of the environment on the survival of a large marine fish population","volume":"85","author":"Ciannelli","year":"2004","journal-title":"Ecology"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"148","DOI":"10.1080\/10705511.2014.935842","article-title":"Structural equation modeling with \u03a9nyx","volume":"22","author":"Brandmaier","year":"2015","journal-title":"Struct. Equ. Model. Multidiscip. J."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"671","DOI":"10.14214\/sf.275","article-title":"Local models for forest canopy cover with beta regression","volume":"41","author":"Korhonen","year":"2007","journal-title":"Silva Fenn."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"e02185","DOI":"10.1002\/ecs2.2185","article-title":"Forests, shrubs, and terrain: Top-down and bottom-up controls on forest structure","volume":"9","author":"Bolstad","year":"2018","journal-title":"Ecosphere"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1093\/forestry\/cpi039","article-title":"Overstorey density influence on the height of Picea abies regeneration in northern Sweden","volume":"78","author":"Chrimes","year":"2005","journal-title":"Forestry"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1016\/j.foreco.2005.12.060","article-title":"Evaluation of relationships between understory light and aspen basal area in the British Columbia central interior","volume":"226","author":"Comeau","year":"2006","journal-title":"For. Ecol. Manag."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"31","DOI":"10.5424\/srf\/2008171-01021","article-title":"Influence of overstory density on understory light, soil moisture, and survival of two underplanted oak species in a Mediterranean montane Scots pine forest","volume":"17","author":"Mutke","year":"2008","journal-title":"For. Syst."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"657","DOI":"10.4067\/S0717-66432018000200657","article-title":"Cambios en cobertura y regeneraci\u00f3n arb\u00f3rea en bosques siempreverdes en diferentes estados sucesionales en el sitio experimental de Llancahue, Cordillera de la Coa de Valdivia, Chile","volume":"75","author":"Donoso","year":"2018","journal-title":"Gayana Bot."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1016\/j.foreco.2017.06.052","article-title":"Importance of structure for species richness and tree species regeneration niches in old-growth Patagonian swamp forests","volume":"401","author":"Bannister","year":"2017","journal-title":"For. Ecol. Manag."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1023\/B:VEGE.0000029336.80225.f0","article-title":"Saplings distribution in relation to topography and canopy openness in an evergreen broad-leaved forest","volume":"173","author":"Enoki","year":"2004","journal-title":"Plant Ecol."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"1438","DOI":"10.1139\/x99-102","article-title":"Evaluation of several measures of canopy openness as predictors of photosynthetic photon flux density in deeply shaded conifer-dominated forest understory","volume":"29","author":"Machado","year":"1999","journal-title":"Can. J. For. Res."},{"key":"ref_76","first-page":"57","article-title":"Canopy openness, understory light environments, and oak regeneration","volume":"Volume 299","author":"Sutherland","year":"2003","journal-title":"Characteristics of Mixed Oak Forest Ecosystems in Southern Ohio Prior to the Reintroduction of Fire"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"118100","DOI":"10.1016\/j.foreco.2020.118100","article-title":"The impact of tree canopy structure on understory variation in a boreal forest","volume":"466","author":"Majasalmi","year":"2020","journal-title":"For. Ecol. Manag."},{"key":"ref_78","unstructured":"Donoso, C. (1993). Estructura, Variaci\u00f3n y Din\u00e1mica de Bosques Templados de Chile y Argentina, Editorial Universitaria. Ecolog\u00eda Forestal."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"1023","DOI":"10.1111\/j.1365-2745.2010.01698.x","article-title":"Differential effects of climate variability on forest dynamics along a precipitation gradient in northern Patagonia","volume":"98","author":"Suarez","year":"2010","journal-title":"J. Ecol."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"119169","DOI":"10.1016\/j.foreco.2021.119169","article-title":"Precipitation declines influence the understory patterns in Nothofagus pumilio old-growth forests in northwestern Patagonia","volume":"491","author":"Soto","year":"2021","journal-title":"For. Ecol. Manag."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1016\/j.rse.2005.08.010","article-title":"Substrate age and precipitation effects on Hawaiian forest canopies from spaceborne imaging spectroscopy","volume":"98","author":"Asner","year":"2005","journal-title":"Remote Sens. Environ."},{"key":"ref_82","unstructured":"Holdridge, R. (1967). Life Zone Ecology, Tropical Science Center."},{"key":"ref_83","unstructured":"Whittaker, R.H. (1970). Communities and Ecosystems, Macmillan."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"727","DOI":"10.1111\/j.1600-0706.2012.20655.x","article-title":"Seasonal, not annual precipitation drives community productivity across ecosystems","volume":"122","author":"Robinson","year":"2013","journal-title":"Oikos"},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"875","DOI":"10.1002\/jgrg.20073","article-title":"The sensitivity of forest water use to the timing of precipitation and snowmelt recharge in the California Sierra: Implications for a warming climate","volume":"118","author":"Tague","year":"2013","journal-title":"J. Geophys. Res. Biogeosci."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"593","DOI":"10.1146\/annurev-ecolsys-110512-135914","article-title":"The structure, distribution, and biomass of the world\u2019s forests","volume":"44","author":"Pan","year":"2013","journal-title":"Annu. Rev. Ecol. Evol. Syst."}],"container-title":["Remote Sensing"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2072-4292\/13\/8\/1513\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T05:48:02Z","timestamp":1760161682000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2072-4292\/13\/8\/1513"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,4,14]]},"references-count":86,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2021,4]]}},"alternative-id":["rs13081513"],"URL":"https:\/\/doi.org\/10.3390\/rs13081513","relation":{},"ISSN":["2072-4292"],"issn-type":[{"value":"2072-4292","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,4,14]]}}}