Dry conditions and disturbance promote liana seedling survival and abundance.

Species composition and community structure in Neotropical forests have been severely affected by increases in climate change and disturbance. Among the most conspicuous changes is the proliferation of lianas. These increases have affected not only the carbon storage capacity of forests but also tree dynamics by reducing tree growth and increasing mortality. Despite the importance of lianas in Neotropical forests, most of the studies on lianas have focused on adult stages, ignoring dynamics at the seedlings stage. Here, we asked whether observed increases in liana abundance are associated with a demographic advantage that emerges early in liana ontogeny and with decreased precipitation and increased disturbance. To test this, we compared patterns of growth and survival between liana seedlings and tree seedlings using a long-term data set of seedling plots from a subtropical wet forest in Puerto Rico, USA. Then, we examined the effect of precipitation and land use history on these demographic variables. We found evidence for liana seedling survival advantage over trees, but no growth advantages. This survival advantage exhibited significant temporal variation linked with patterns of rainfall, as well as differences associated with land-use history in the study area. Furthermore, we found that neighborhood density has a negative effect on liana survival and growth. Our results indicate that liana proliferation is likely related to a survival advantage that emerges in early stages and is influenced by climatic conditions and past disturbance. Predicted climatic changes in rainfall patterns, including more frequent and severe droughts, together with increases in disturbance, could have a significant effect on seedling tropical communities by favoring lianas.

[1]  F. Bazzaz,et al.  13 – Coping with Environmental Heterogeneity: The Physiological Ecology of Tree Seedling Regeneration across the Gap—Understory Continuum , 1994 .

[2]  J. Zimmerman,et al.  Life‐history trade‐offs during the seed‐to‐seedling transition in a subtropical wet forest community , 2013 .

[3]  Y. Masrahi Ecological significance of wood anatomy in two lianas from arid southwestern Saudi Arabia. , 2014, Saudi journal of biological sciences.

[4]  A. Packer,et al.  SOIL PATHOGENS AND PRUNUS SEROTINA SEEDLING AND SAPLING GROWTH NEAR CONSPECIFIC TREES , 2003 .

[5]  H.,et al.  1 The distribution and evolution of climbing plants , 2012 .

[6]  S. Schnitzer,et al.  Unique competitive effects of lianas and trees in a tropical forest understory , 2015, Oecologia.

[7]  P. Fearnside,et al.  RAIN FOREST FRAGMENTATION AND THE STRUCTURE OF AMAZONIAN LIANA COMMUNITIES , 2001 .

[8]  D. Nepstad,et al.  Seedling growth dynamics of a deeply rooting liana in a secondary forest in eastern Amazonia , 2004 .

[9]  Bettina M. J. Engelbrecht,et al.  Forests and Global Change: Drought as a driver of tropical tree species regeneration dynamics and distribution patterns , 2014 .

[10]  Benjamin L Turner,et al.  Liana effects on biomass dynamics strengthen during secondary forest succession. , 2017, Ecology.

[11]  N. McDowell,et al.  A global overview of drought and heat-induced tree mortality reveals emerging climate change risks for forests , 2010 .

[12]  F. Bongers,et al.  Contrasting nitrogen and phosphorus resorption efficiencies in trees and lianas from a tropical montane rain forest in Xishuangbanna, south-west China , 2007, Journal of Tropical Ecology.

[13]  R. Valencia,et al.  No strong evidence for increasing liana abundance in the Myristicaceae of a Neotropical aseasonal rain forest. , 2017, Ecology.

[14]  C. Vriesendorp,et al.  Conspecific density dependence in seedlings varies with species shade tolerance in a wet tropical forest. , 2011, Ecology letters.

[15]  Susan G. Letcher,et al.  Lianas and self-supporting plants during tropical forest succession. , 2009 .

[16]  Charles D. Canham,et al.  Natural disturbance and human land use as determinants of tropical forest dynamics: results from a forest simulator , 2009 .

[17]  N. Brokaw,et al.  Forest structure before and after Hurricane Hugo at three elevations in the Luquillo Mountains, Puerto Rico , 1991 .

[18]  S. Hubbell,et al.  The impact of lianas on 10 years of tree growth and mortality on Barro Colorado Island, Panama , 2010 .

[19]  W. Carson,et al.  Lianas suppress tree regeneration and diversity in treefall gaps. , 2010, Ecology letters.

[20]  P. Woods,et al.  Effects of Logging, Drought, and Fire on Structure and Composition of Tropical Forests in Sabah, Malaysia , 1989 .

[21]  W. Carson,et al.  Drought stress and tropical forest woody seedlings: effect on community structure and composition , 2005 .

[22]  F. Putz Liana biomass and leaf area of a «Tierra Firme» forest in the Rio Negro Basin, Venezuela , 1983 .

[23]  Stephen P. Hubbell,et al.  Drought sensitivity shapes species distribution patterns in tropical forests , 2007, Nature.

[24]  O. Phillips,et al.  Increasing Turnover Through Time in Tropical Forests , 1994, Science.

[25]  Frans Bongers,et al.  Above-ground biomass and productivity in a rain forest of eastern South America , 2008, Journal of Tropical Ecology.

[26]  S. Schnitzer A Mechanistic Explanation for Global Patterns of Liana Abundance and Distribution , 2005, The American Naturalist.

[27]  Kyle G. Dexter,et al.  Seasonal drought limits tree species across the Neotropics , 2017 .

[28]  G. V. D. van der Heijden,et al.  Lianas reduce carbon accumulation and storage in tropical forests , 2015, Proceedings of the National Academy of Sciences.

[29]  D. Bates,et al.  Fitting Linear Mixed-Effects Models Using lme4 , 2014, 1406.5823.

[30]  Robert B. Waide,et al.  Responses of Tree Species to Hurricane Winds in Subtropical Wet Forest in Puerto Rico: Implications for Tropical Tree Life Histories , 1994 .

[31]  S. Schnitzer,et al.  Lianas Have a Greater Competitive Effect Than Trees of Similar Biomass on Tropical Canopy Trees , 2012 .

[32]  R. Ong,et al.  Intra‐annual plasticity of growth mediates drought resilience over multiple years in tropical seedling communities , 2017, Global change biology.

[33]  Emery R. Boose,et al.  HUMAN OR NATURAL DISTURBANCE: LANDSCAPE‐SCALE DYNAMICS OF THE TROPICAL FORESTS OF PUERTO RICO , 1999 .

[34]  F. Putz,et al.  Lianas and Trees in a Liana Forest of Amazonian Bolivia1 , 2001 .

[35]  Frank W. Ewers,et al.  Xylem' Structure and Water Conduction in Conifer Trees, Dicot Trees, and Llanas , 1985 .

[36]  Robert B. Waide,et al.  Land use history, environment, and tree composition in a tropical forest , 2002 .

[37]  S. Vieira,et al.  Variation in liana abundance and biomass along an elevational gradient in the tropical Atlantic Forest (Brazil) , 2012, Ecological Research.

[38]  A. Gentry The Biology of Vines : The distribution and evolution of climbing plants , 1992 .

[39]  Yadvinder Malhi,et al.  Increasing dominance of large lianas in Amazonian forests , 2002, Nature.

[40]  Noelle G. Beckman,et al.  Testing predictions of the Janzen–Connell hypothesis: a meta-analysis of experimental evidence for distance- and density-dependent seed and seedling survival , 2014, The Journal of ecology.

[41]  J. Fisher,et al.  A survey of vessel dimensions in stems of tropical lianas and other growth forms , 1990, Oecologia.

[42]  Brian J. Enquist,et al.  Long‐term change within a Neotropical forest: assessing differential functional and floristic responses to disturbance and drought , 2011 .

[43]  F. Putz,et al.  Ecology of Lianas , 2014 .

[44]  J. Zimmerman,et al.  Environmental heterogeneity and biotic interactions mediate climate impacts on tropical forest regeneration , 2018, Global change biology.

[45]  G. C. Stevens Lianas as structural parasites: the Bursera simaruba example , 1987 .

[46]  Stefan A. Schnitzer,et al.  Density and diversity of lianas along a chronosequence in a central Panamanian lowland forest , 2000, Journal of Tropical Ecology.

[47]  J. Zimmerman,et al.  Liana dynamics reflect land-use history and hurricane response in a Puerto Rican forest , 2017, Journal of Tropical Ecology.

[48]  J. Grace,et al.  Lianas may be favoured by low rainfall: evidence from Ghana , 2007, Plant Ecology.

[49]  I. Sun,et al.  Long-term changes in liana loads and tree dynamics in a Malaysian forest. , 2015, Ecology.

[50]  F. Bongers,et al.  Increasing liana abundance and biomass in tropical forests: emerging patterns and putative mechanisms. , 2011, Ecology letters.

[51]  P. Reich,et al.  Species with greater seed mass are more tolerant of conspecific neighbours: a key driver of early survival and future abundances in a tropical forest. , 2016, Ecology letters.

[52]  R. Condit Ecological Implications of Changes in Drought Patterns: Shifts in Forest Composition in Panama , 1998 .

[53]  F. Bongers,et al.  Seasonal differences in leaf-level physiology give lianas a competitive advantage over trees in a tropical seasonal forest , 2009, Oecologia.

[54]  R. Nemani,et al.  Persistent effects of a severe drought on Amazonian forest canopy , 2012, Proceedings of the National Academy of Sciences.

[55]  V. Angyalossy,et al.  Liana anatomy: a broad perspective on structural evolution of the vascular system , 2014 .

[56]  D. Nepstad,et al.  Mortality of large trees and lianas following experimental drought in an Amazon forest. , 2007, Ecology.

[57]  Frans Bongers,et al.  The ecology of lianas and their role in forests , 2002 .

[58]  James C. Stegen,et al.  Disentangling the Drivers of β Diversity Along Latitudinal and Elevational Gradients , 2011, Science.

[59]  S. Paton,et al.  ARE LIANAS INCREASING IN IMPORTANCE IN TROPICAL FORESTS? A 17‐YEAR RECORD FROM PANAMA , 2004 .

[60]  C. Restrepo,et al.  Using multiple traits to assess the potential of introduced and native vines to proliferate in a tropical region , 2016, Ecology and evolution.