Demographic trade-offs and functional shifts in a hurricane-impacted tropical forest.

BACKGROUND AND AIMS: Understanding shifts in the demographic and functional composition of forests after major natural disturbances has become increasingly relevant given the accelerating rates of climate change and elevated frequency of natural disturbances. Although plant demographic strategies are often described across a slow-fast continuum, severe and frequent disturbance events influencing demographic processes may alter the demographic trade-offs and the functional composition of forests. We examined demographic trade-offs and the shifts in functional traits in a hurricane-disturbed forest using long-term data from the Luquillo Forest Dynamics Plot (LFPD) in Puerto Rico. METHODS: We analysed information on growth, survival, seed rain and seedling recruitment for 30 woody species in the LFDP. In addition, we compiled data on leaf, seed and wood functional traits that capture the main ecological strategies for plants. We used this information to identify the main axes of demographic variation for this forest community and evaluate shifts in community-weighted means for traits from 2000 to 2016. KEY RESULTS: The previously identified growth-survival trade-off was not observed. Instead, we identified a fecundity-growth trade-off and an axis representing seedling-to-adult survival. Both axes formed dimensions independent of resprouting ability. Also, changes in tree species composition during the post-hurricane period reflected a directional shift from seedling and tree communities dominated by acquisitive towards conservative leaf economics traits and large seed mass. Wood specific gravity, however, did not show significant directional changes over time. CONCLUSIONS: Our study demonstrates that tree demographic strategies coping with frequent storms and hurricane disturbances deviate from strategies typically observed in undisturbed forests, yet the shifts in functional composition still conform to the expected changes from acquisitive to conservative resource-uptake strategies expected over succession. In the face of increased rates of natural and anthropogenic disturbance in tropical regions, our results anticipate shifts in species demographic trade-offs and different functional dimensions.

Proposing the solar-wind energy flux hypothesis as a driver of inter-annual variation in tropical tree reproductive effort.

PREMISE: The El Niño Southern Oscillation (ENSO) affects tropical environmental conditions, potentially altering ecosystem function as El Niño events interact with longer-term climate change. Anomalously warm equatorial Pacific Ocean temperatures affect rainfall and temperature throughout the tropics and coincide with altered leaf flush phenology and increased fruit production in wet tropical forests; however, the understanding of mechanisms underlying this pattern is limited. There is evidence that increases in tropical tree reproduction anticipate El Niño onset, motivating the continued search for a global driver of tropical angiosperm reproduction. We present the solar-wind energy flux hypothesis: that physical energy influx to the Earth's upper atmosphere and magnetosphere, generated by a positive anomaly in the solar wind preceding El Niño development, cues tropical trees to increase resource allocation to reproduction. METHODS: We test this hypothesis using 19 years of data from Luquillo, Puerto Rico, correlating them with measures of solar-wind energy. RESULTS: From 1994 to 2013, the solar-wind energy flux into Earth's magnetosphere (Ein ) was more strongly correlated with the number of species fruiting and flowering than the Niño 3.4 climate index, despite Niño 3.4 being previously identified as a driver of interannual increases in reproduction. CONCLUSIONS: Changes in the global magnetosphere and thermosphere conditions from increased solar-wind energy affect global atmospheric pressure and circulation patterns, principally by weakening the Walker circulation. We discuss the idea that these changes cue interannual increases in tropical tree reproduction and act through an unidentified mechanism that anticipates and synchronizes the reproductive output of the tropical trees with El Niño.

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.

Effects of hurricanes and climate oscillations on annual variation in reproduction in wet forest, Puerto Rico.

Interannual changes in global climate and weather disturbances may influence reproduction in tropical forests. Phenomena such as the El Niño Southern Oscillation (ENSO) are known to produce interannual variation in reproduction, as do severe storms such as hurricanes. Using stationary trap-based phenology data collected fortnightly from 1993 to 2014 from a hurricane-affected (1989 Hugo, 1998 Georges) subtropical wet forest in northeastern Puerto Rico, we conducted a time series analysis of flowering and seed production. We addressed (1) the degree to which interannual variation in flower and seed production was influenced by global climate drivers and time since hurricane disturbance, and (2) how long-term trends in reproduction varied with plant lifeform. The seasonally de-trended number of species in flower fluctuated over time while the number of species producing seed exhibited a declining trend, one that was particularly evident during the second half of the study period. Lagged El Niño indices and time series hurricane disturbance jointly influenced the trends in numbers of flowering and fruiting species, suggesting complex global influences on tropical forest reproduction with variable periodicities. Lag times affecting flowering tended to be longer than those affecting fruiting. Long-term patterns of reproduction in individual lifeforms paralleled the community-wide patterns, with most groups of lifeform exhibiting a long-term decline in seed but not flower production. Exceptions were found for hemiepiphytes, small trees, and lianas whose seed reproduction increased and then declined over time. There was no long-term increase in flower production as reported in other Neotropical sites.

Interspecific Functional Convergence and Divergence and Intraspecific Negative Density Dependence Underlie the Seed-to-Seedling Transition in Tropical Trees.

The seed-to-seedling transition constitutes a critical bottleneck in the life history of plants and represents a major determinant of species composition and abundance. However, we have surprisingly little knowledge regarding the forces driving this ontogenetic transition. Here we utilize information regarding organismal function to investigate the strength of intra- and interspecific negative density dependence during the seed-to-seedling transition in Puerto Rican tree species. Our analyses were implemented at individual sites and across an entire 16-ha forest plot, spanning 6 years. The functional richness of seedling assemblages was significantly lower than expected given the seed assemblages, but the functional evenness was significantly higher than expected, indicating the simultaneous importance of constraints on the overall phenotypic space and trait differences for successful transitions from seed to seedling. The results were consistent across years. Within species, we also found evidence for strong intraspecific negative density dependence, where the probability of transition was proportionally lower when in a site with high conspecific density. These results suggest that filtering of similar phenotypes across species and strong negative density dependence within and among species are simultaneously driving the structure and dynamics of tropical tree assemblages during this critical life-history transition.

Ontogenetic shifts in trait-mediated mechanisms of plant community assembly.

Identifying the processes that maintain highly diverse plant communities remains a central goal in ecology. Species variation in growth and survival rates across ontogeny, represented by tree size classes and life history stage-specific niche partitioning, are potentially important mechanisms for promoting forest diversity. However, the role of ontogeny in mediating competitive dynamics and promoting functional diversity is not well understood, particular in high-diversity systems such as tropical forests. The interaction between interspecific functional trait variation and ontogenetic shifts in competitive dynamics may yield insights into the ecophysiological mechanisms promoting community diversity. We investigated how functional trait (seed size, maximum height, SLA, leaf N, and wood density) associations with growth, survival, and response to competing neighbors differ among seedlings and two size classes of trees in a subtropical rain forest in Puerto Rico. We used a hierarchical Bayes model of diameter growth and survival to infer trait relationships with ontogenetic change in competitive dynamics. Traits were more strongly associated with average growth and survival than with neighborhood interactions, and were highly consistent across ontogeny for most traits. The associations between trait values and tree responses to crowding by neighbors showed significant shifts as trees grew. Large trees exhibited greater growth as the difference in species trait values among neighbors increased, suggesting trait-associated niche partitioning was important for the largest size class. Our results identify potential axes of niche partitioning and performance-equalizing functional trade-offs across ontogeny, promoting species coexistence in this diverse forest community.