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Summary: Among tropical forests, lianas are predicted to have a growth advantage over trees during seasonal drought, with substantial implications for tree and forest dynamics. We tested the hypotheses that lianas maintain higher water status than trees during seasonal drought and that lianas maximize leaf cover to match high, dry‐season light conditions, while trees are more limited by moisture availability during the dry season.We monitored the seasonal dynamics of predawn and midday leaf water potentials and leaf phenology for branches of 16 liana and 16 tree species in the canopies of two lowland tropical forests with contrasting rainfall regimes in Panama.In a wet, weakly seasonal forest, lianas maintained higher water balance than trees and maximized their leaf cover during dry‐season conditions, when light availability was high, while trees experienced drought stress. In a drier, strongly seasonal forest, lianas and trees displayed similar dry season reductions in leaf cover following strong decreases in soil water availability.Greater soil moisture availability and a higher capacity to maintain water status allow lianas to maintain the turgor potentials that are critical for plant growth in a wet and weakly seasonal forest but not in a dry and strongly seasonal forest. [ABSTRACT FROM AUTHOR]

Questions remain as to which soil nutrients limit primary production in tropical forests. Phosphorus (P) has long been considered the primary limiting element in lowland forests, but recent evidence demonstrates substantial heterogeneity in response to nutrient addition, highlighting a need to understand and diagnose nutrient limitation across diverse forests. Fine-root characteristics including their abundance, functional traits, and mycorrhizal symbionts can be highly responsive to changes in soil nutrients and may help to diagnose nutrient limitation. Here, we document the response of fine roots to long-term nitrogen (N), P, and potassium (K) fertilization in a lowland forest in Panama. Because this experiment has demonstrated that N and K together limit tree growth and P limits fine litter production, we hypothesized that fine roots would also respond to nutrient addition. Specifically we hypothesized that N, P, and K addition would reduce the biomass, diameter, tissue density, and mycorrhizal colonization of fine roots, and increase nutrient concentration in root tissue. Most morphological root traits responded to the single addition of K and the paired addition of N and P, with the greatest response to all three nutrients combined. The addition of N, P, and K together reduced fine-root biomass, length, and tissue density, and increased specific root length, whereas root diameter remained unchanged. Nitrogen addition did not alter root N concentration, but P and K addition increased root P and K concentration, respectively. Mycorrhizal colonization of fine roots declined with N, increased with P, and was unresponsive to K addition. Although plant species composition remains unchanged after 14 years of fertilization, fine-root characteristics responded to N, P, and K addition, providing some of the strongest stand-level responses in this experiment. Multiple soil nutrients regulate fine-root abundance, morphological and chemical traits, and their association with mycorrhizal fungi in a species-rich lowland tropical forest. [ABSTRACT FROM AUTHOR]

Defaunation alters trophic interactions between plants and vertebrates, which may disrupt trophic cascades, thereby favoring a subset of plant species and reducing diversity. If particular functional traits characterize the favored plant species, then defaunation may alter community-wide patterns of functional trait composition. Changes in plant functional traits occurring with defaunation may help identify the species interactions affected by defaunation and the potential for other cascading effects of defaunation. We tested the hypotheses that defaunation would (1) disrupt seed dispersal, thereby favoring species whose dispersal agents are not affected (e.g., small birds, bats, and abiotic agents), (2) reduce seed predation, thereby favoring larger-seeded species, and (3) reduce herbivory, thereby favoring species with lower leaf mass per area (LMA), leaf toughness, and wood density. We examined how these six traits responded to vertebrate defaunation caused by hunters or by experimental exclosures among more than 30 000 woody seedlings in a lowland tropical moist forest. Exclosures reduced terrestrial frugivores, granivores, and herbivores, while hunters also reduced volant and arboreal frugivores and granivores. The comparison of exclosures and hunting allowed us to parse the impacts of arboreal and volant species (reduced by hunters only) and terrestrial species (reduced by both hunters and exclosures). The loss of terrestrial vertebrates alone had limited effects on plant trait composition. The additional loss of volant and arboreal vertebrates caused significant shifts in plant species composition towards communities with more species dispersed abiotically, including lianas and low wood-density tree species, and fewer species dispersed by large vertebrates. In contrast to previous studies, community seed mass did not decline significantly in hunted sites. Our exclosure results suggest this is because reducing seed predators disproportionately benefits large-seeded species, partially compensating for the reduction of seed dispersers at hunted sites. Our results demonstrate that decreased seed dispersal and seed predation are important determinants of seedling community compositional change as a consequence of defaunation. Defaunation may also negatively impact carbon storage in tropical forests, by favoring lianas and low wood density tree species. [ABSTRACT FROM AUTHOR]

Over the past two decades, liana density and basal area have been increasing in many tropical forests, which has profound consequences for forest diversity and functioning. One hypothesis to explain increasing lianas is elevated nutrient deposition in tropical forests resulting from fossil fuels, agricultural fertilizer, and biomass burning. We tested this hypothesis by surveying all lianas ≥1 cm in diameter (n = 3,967) in 32 plots in a fully factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment in a mature tropical forest in central Panama. We conducted the nutrient‐addition experiment from 1998 until present and we first censused lianas in 2013 and then again in 2018. After 20 yr of nutrient addition (1998–2018), liana density, basal area, and rarefied species richness did not differ significantly among any of the nutrient‐addition and control treatments. Moreover, nutrient addition in the most recent 5 yr of the experiment did not affect liana relative growth, recruitment, or mortality rates. From 2013 until 2018, liana density, basal area, and species richness increased annually by 1.6%, 1.4%, and 2.4%, respectively. Nutrient addition did not influence these increases. Our findings indicate that nutrient deposition does not explain increasing lianas in this tropical forest. Instead, increases in tree mortality and disturbance, atmospheric carbon dioxide, drought frequency and severity, and hunting pressure may be more likely explanations for the increase in lianas in tropical forests. [ABSTRACT FROM AUTHOR]

As extreme climate events are predicted to become more frequent because of global climate change, understanding their impacts on natural systems is crucial. Tropical forests are vulnerable to droughts associated with extreme El Niño events. However, little is known about how tropical seedling communities respond to El Niño–related droughts, even though patterns of seedling survival shape future forest structure and diversity. Using long‐term data from eight tropical moist forests spanning a rainfall gradient in central Panama, we show that community‐wide seedling mortality increased by 11% during the extreme 2015–16 El Niño, with mortality increasing most in drought‐sensitive species and in wetter forests. These results indicate that severe El Niño–related droughts influence understory dynamics in tropical forests, with effects varying both within and across sites. Our findings suggest that predicted increases in the frequency of extreme El Niño events will alter tropical plant communities through their effects on early life stages. [ABSTRACT FROM AUTHOR]

The top‐down and indirect effects of insects on plant communities depend on patterns of host use, which are often poorly documented, particularly in species‐rich tropical forests. At Barro Colorado Island, Panama, we compiled the first food web quantifying trophic interactions between the majority of co‐occurring woody plant species and their internally feeding insect seed predators. Our study is based on more than 200 000 fruits representing 478 plant species, associated with 369 insect species. Insect host‐specificity was remarkably high: only 20% of seed predator species were associated with more than one plant species, while each tree species experienced seed predation from a median of two insect species. Phylogeny, but not plant traits, explained patterns of seed predator attack. These data suggest that seed predators are unlikely to mediate indirect interactions such as apparent competition between plant species, but are consistent with their proposed contribution to maintaining plant diversity via the Janzen–Connell mechanism. [ABSTRACT FROM AUTHOR]

Myxomycetes (plasmodial slime molds) are abundant protist predators that feed on bacteria and other microorganisms, thereby playing important roles in terrestrial nutrient cycling. Despite their significance, little is known about myxomycete communities and the extent to which they are affected by nutrient availability. We studied the influence of long‐term addition of N, P, and K on the myxomycete community in a lowland forest in the Republic of Panama. In a previous study, microbial biomass increased with P but not N or K addition at this site. We hypothesized that myxomycetes would increase in abundance in response to P but that they would not respond to the sole addition of N or K. Moist chamber cultures of leaf litter and small woody debris were used to quantify myxomycete abundance. We generated the largest myxomycete dataset (3,381 records) for any single locality in the tropics comprised by 91 morphospecies. In line with our hypothesis, myxomycete abundance increased in response to P addition but did not respond to N or K. Community composition was unaffected by nutrient treatments. This work represents one of very few large‐scale and long‐term field studies to include a heterotrophic protist highlighting the feasibility and value in doing so. [ABSTRACT FROM AUTHOR]

Nonstructural carbohydrates (NSCs) are essential for maintenance of plant metabolism and may be sensitive to short‐ and long‐term climatic variation. NSC variation in moist tropical forests has rarely been studied, so regulation of NSCs in these systems is poorly understood. We measured foliar and branch NSC content in 23 tree species at three sites located across a large precipitation gradient in Panama during the 2015–2016 El Niño to examine how short‐ and long‐term climatic variation impact carbohydrate dynamics. There was no significant difference in total NSCs as the drought progressed (leaf P = 0.32, branch P = 0.30) nor across the rainfall gradient (leaf P = 0.91, branch P = 0.96). Foliar soluble sugars decreased while starch increased over the duration of the dry period, suggesting greater partitioning of NSCs to storage than metabolism or transport as drought progressed. There was a large variation across species at all sites, but total foliar NSCs were positively correlated with leaf mass per area, whereas branch sugars were positively related to leaf temperature and negatively correlated with daily photosynthesis and wood density. The NSC homoeostasis across a wide range of conditions suggests that NSCs are an allocation priority in moist tropical forests. Nonstructural carbohydrates (NSCs), the organic compounds that drive plant metabolism, have rarely been studied in moist tropical forests, so their regulation in these systems is poorly understood. We measured foliar and branch NSCs in canopy trees across a large precipitation gradient in Panama during the 2015–2016 El Niño to examine how short‐ and long‐term climatic variation impact carbohydrate dynamics. There was a large variation in NSCs across species; however, there was no change in total NSCs as the drought progressed nor across the rainfall gradient, suggesting that NSCs are an allocation priority in moist tropical forests. [ABSTRACT FROM AUTHOR]

Local tree species distributions in tropical forests correlate strongly with soil water availability. However, it is unclear how species distributions are shaped by demographic responses to soil water availability. Specifically, it remains unknown how growth affects species distributions along water availability gradients relative to mortality.We quantified spatial variation in dry season soil water potential (SWP) in the moist tropical forest on Barro Colorado Island, Panama, and used a hierarchical Bayesian approach to evaluate relationships between demographic responses of naturally regenerating seedlings to SWP (RGRs and first‐year mortality) and species distributions along the SWP gradient for 62 species. We also tested whether species that were more abundant at the wet or dry end of the gradient performed better (a) at their "home end" of the gradient ("best at home" hypothesis) and (b) "at home" compared to co‐occurring species ("home advantage" hypothesis).Four and five species responded significantly to SWP in terms of growth or mortality respectively. Growth (but not mortality) responses were positively related to species distributions along the SWP gradient; species with a more positive (negative) growth response to SWP were more abundant at higher (lower) SWP, that is, at wetter (drier) sites. In addition, wet distributed species grew faster on the wet end of the SWP gradient than on the dry end ("best at home") and grew faster on the wet end than dry distributed species ("home advantage"). Mortality rates declined with seedling size for all species. Thus, seedling growth responses to SWP indirectly shaped local species distributions by influencing seedling size and thereby mortality risk.Synthesis. By demonstrating how growth responses to spatial variation in soil water availability affect species distributions, we identified a demographic process underlying niche differentiation on hydrological gradients in tropical forests. Recognizing the role of these growth responses in shaping species distributions should improve the understanding of tropical forest composition and diversity along rainfall gradients and with climate change. This study shows that growth responses to soil moisture play an important role in shaping the distribution of tree seedlings along a local soil moisture gradient. Species that grew well on the wet end of the gradient were more abundant there because they became taller more quickly, reducing their mortality risk, and because they grew faster than dry distributed species. [ABSTRACT FROM AUTHOR]

Abstract: Lianas are structural parasites of trees and reduce individual host tree growth, survival and fecundity. Thus, liana infestation is expected to affect tree population growth rates, with potentially different effects in different species depending on the frequency of liana infestation and the impact of liana infestation on population growth rates. Previous studies have documented the myriad negative effects of lianas on trees and variation in liana infestation among tree species; however, no study has quantified the impact of liana infestation on individual tree species population growth rates. Lianas are increasing in abundance in multiple Neotropical sites, which may have profound consequences for tree species composition if lianas differentially affect host tree species population growth. Here, we use long‐term data to evaluate the effects of liana infestation on the reproduction, growth, survival and ultimately population growth rates of dozens of tree species from Barro Colorado Island, Panama. We then test whether liana infestation affects tree species differentially with respect to two axes of life‐history variation: adult stature and position along the fast–slow axis, a measure of shade tolerance. Liana infestation decreased tree growth, survival and reproduction, with the strongest effects on survival in fast‐growing, light‐demanding species and on reproduction in large‐statured species. In combination, these effects reduced tree population growth rates such that liana‐infested populations declined by an average of 1.4% annually relative to conspecific liana‐free populations. The reduction in population growth rates was greatest among fast‐growing species and smaller in slow‐growing species. Synthesis. Our results demonstrate that liana infestation has strong negative effects on tree population growth rates, which vary systematically among tree species with tree life history. The finding that liana infestation is more harmful to fast‐growing tree species appears to be at odds with the general expectations in the literature. We propose that this is likely due to survivorship bias, as infestation greatly decreases survival in fast‐growing species such that the observable sample is biased towards those that survived and liana‐free. In combination with data on how tree species vary in liana infestation rates, these results provide a basis for predicting the impacts of changes in liana abundance on tree species composition. [ABSTRACT FROM AUTHOR]

1. Identification of the mechanisms enabling stable coexistence of species with similar resource requirements is a central challenge in ecology. Such coexistence can be facilitated by species at higher trophic levels through complex multi-trophic interactions, a mechanism that could be compromised by ongoing defaunation. 2. We investigated cascading effects of defaunation on Pachymerus cardo and Speciomerus giganteus, the specialized insect seed predators of the Neotropical palm Attalea butyracea, testing the hypothesis that vertebrate frugivores and granivores facilitate their coexistence. 3. Laboratory experiments showed that the two seed parasitoid species differed strongly in their reproductive ecology. Pachymerus produced many small eggs that it deposited exclusively on the fruit exocarp (exterior). Speciomerus produced few large eggs that it deposited exclusively on the endocarp, which is normally exposed only after a vertebrate handles the fruit. When eggs of the two species were deposited on the same fruit, Pachymerus triumphed only when it had a long head start, and the loser always succumbed to intraguild predation. 4. We collected field data on the fates of 6569 Attalea seeds across sites in central Panama with contrasting degrees of defaunation and wide variation in the abundance of vertebrate frugivores and granivores. Speciomerus dominated where vertebrate communities were intact, whereas Pachymerus dominated in defaunated sites. Variation in the relative abundance of Speciomerus across all 84 sampling sites was strongly positively related to the proportion of seeds attacked by rodents, an indicator of local vertebrate abundance. 5. Synthesis. We show that two species of insect seed predators relying on the same host plant species are niche differentiated in their reproductive strategies such that one species has the advantage when fruits are handled promptly by vertebrates and the other when they are not. Defaunation disrupts this mediating influence of vertebrates and strongly favours one species at the expense of the other, providing a case study of the cascading effects of defaunation and its potential to disrupt coexistence of non-target species, including the hyperdiverse phytophagous insects of tropical forests. [ABSTRACT FROM AUTHOR]

Soil organic matter is an important pool of carbon and nutrients in tropical forests. The majority of this pool is assumed to be relatively stable and to turn over slowly over decades to centuries, although changes in nutrient status can influence soil organic matter on shorter timescales. We measured carbon, nitrogen, and phosphorus concentrations in soil organic matter and leaf litter over an annual cycle in a long-term nutrient addition experiment in lowland tropical rain forest in the Republic of Panama. Total soil carbon was not affected by a decade of factorial combinations of nitrogen, phosphorus, or potassium. Nitrogen addition increased leaf litter nitrogen concentration by 7 % but did not affect total soil nitrogen. Phosphorus addition doubled the leaf litter phosphorus concentration and increased soil organic phosphorus by 50 %. Surprisingly, concentrations of carbon, nitrogen, and phosphorus in soil organic matter declined markedly during the four-month dry season, and then recovered rapidly during the following wet season. Between the end of the wet season and the late dry season, total soil carbon declined by 16 %, total nitrogen by 9 %, and organic phosphorus by between 19 % in control plots and 25 % in phosphorus addition plots. The decline in carbon and nitrogen was too great to be explained by changes in litter fall, bulk density, or the soil microbial biomass. However, a major proportion of the dry-season decline in soil organic phosphorus was explained by a corresponding decline in the soil microbial biomass. These results have important implications for our understanding of the stability and turnover of organic matter in tropical forest soils, because they demonstrate that a considerable fraction of the soil organic matter is seasonally transient, despite the overall pool being relatively insensitive to long-term changes in nutrient status. [ABSTRACT FROM AUTHOR]