The invasive tropical shrub Clidemia hirta (Melastomataceae) in its native and introduced ranges: tests of hypotheses of invasion

Exotic pest plants often grow to greater stature, become more abundant, and display increased shade tolerance in their introduced ranges than in their native ranges. These differences have been hypothesized to result from genetic shifts in biomass allocation, growth, or photosynthesis between genotypes in native and introduced ranges or from plastic, phenotypic responses to different environmental conditions, such as lower herbivore or fungal pest loads in areas of introduction. I used the tropical shrub Clidemia hirta (Melastomataceae) as a model exotic pest plant to test these two non-mutually exclusive hypotheses of invasion. Clidemia hirta invades forest understory and is more abundant in much of its introduced range in parts of Oceania, Asia, and Africa than in its native range in Central and South America, where it does not occur in forest understory. Contrary to predictions, I found less genetic variation, as detected with allozymes, within and among native, Costa Rican populations than introduced, Hawaiian populations of C. hirta. Hawaiian and Costa Rican populations also were markedly dissimilar genetically (Nei\u27s I = 0.64), but there were few ecologically important differences in biomass allocation, growth, or photosynthetic parameters between Costa Rican and Hawaiian genotypes grown under high or low light in a common garden experiment. The absence of C. hirta from forest understory in its native range likely results, at least in part, from the strong pressures of insect herbivores and pathogens (natural enemies). A natural enemy exclusion study conducted in the field showed that insect herbivore and fungal pathogen damage was substantially greater on Costa Rican than Hawaiian plants and that these natural enemies caused substantial mortality of C. hirta planted into forest understory in Costa Rica but not Hawaii. These results coupled with demographic data collected over three years in two Hawaiian populations suggest that biological control could cause a decline in C. hirta population growth rates in Hawaiian forests. For now the expanded habitat distribution and vigor of C. hirta in its introduced range seems to result from an ecological response to enemy release rather than a genetic shift in resource acquisition, allocation, or growth

Censusing and Measuring Lianas: A Quantitative Comparison of the Common Methods

Lianas contribute to many aspects of tropical forest diversity and dynamics, and interest in liana ecology has grown substantially in recent years. Methods to census lianas and estimate biomass, however, differ among studies, possibly hindering attempts to compare liana communities. At Nouragues Research Station (French Guiana), we tested the extent to which liana abundance, basal area, and estimated biomass differed depending on stem diameter measurement location, inclusion of ramets, inclusion of lianas rooted within versus passing through the plot, and plot shape. We found that the mean per plot abundance and basal area of lianas were significantly greater when lianas were measured low on the stem, when ramets were included, and when lianas were sampled in transects (2 × 50 m) than in square plots (10 × 10 m). Mean per plot liana abundance and basal area were 21 percent and 58 percent greater, when stems were measured at the largest spot on the stem compared to 130 cm from the ground, respectively. Including liana ramets increased average per plot liana abundance, basal area, and estimated biomass by 19, 17, and 16 percent, respectively. To facilitate cross‐study comparisons, we developed conversion equations that equate liana abundance, diameter, and basal area based on the measurements taken at four different stem locations. We tested these equations at Lambir Hills National Park, Malaysia and found that they did not differ significantly between the two sites, suggesting that the equations may be broadly applicable. Finally, we present a new allometric equation relating diameter and biomass developed from 424 lianas from five independent data sets collected in four countries

Liana habitat associations and community structure in a Bornean lowland tropical forest

Lianas (woody vines) contribute substantially to the diversity and structure of most tropical forests, yet little is known about the importance of habitat specialization in maintaining tropical liana diversity and the causes of variation among forests in liana abundance and species composition. We examined habitat associations, species diversity, species composition, and community structure of lianas at Sepilok Forest Reserve, Sabah, Malaysia in northeastern Borneo among three soil types that give rise to three distinct forest types of lowland tropical rain forest: alluvial, sandstone hill, and kerangas (heath) forest. Alluvial soils are more nutrient rich and have higher soil moisture than sandstone soils, whereas kerangas soils are the most nutrient poor and drought prone. Lianas ≥0.5-cm in diameter were measured, tagged, and identified to species in three square 0.25-ha plots in each forest type. The number of lianas ≥0.5 cm did not differ significantly among forest types and averaged 1348 lianas ha-1, but mean liana stem diameter, basal area, estimated biomass, species richness, and Fisher\u27s α diversity index were all greater for plots in alluvial than sandstone or kerangas forests. Liana species composition also differed greatly among the three habitats, with 71% of species showing significant positive or negative habitat associations. Sandstone forests were intermediate to alluvial and kerangas forests in most aspects of liana community structure and composition, and fewer species showed significant habitat associations with this forest type. Ranking of forest types with respect to liana density, biomass, and diversity matches the ranking in soil fertility and water availability (alluvial \u3e sandstone hill \u3e kerangas). These results suggest that edaphic factors play an important role in maintaining liana species diversity and structuring liana communities. © Springer 2006

Fertilization influences the nutrient acquisition strategy of a nomadic vine in a lowland tropical forest understory

© 2018, Springer Nature Switzerland AG. Aims: Tropical tree and lianas in the understory are limited by soil nutrients despite growing in extremely low light. It is not known if nomadic vines are also limited by nutrients in low light conditions. Methods: We measured differences in root architecture and mycorrhizal colonization, and leaf nutrients of a nomadic vine, Philodendron fragrantissimum (Araceae), in nitrogen (N) and phosphorus (P) fertilization plots in a lowland tropical moist forest in central Panama to measure potential nutrient limitation. Results: Relative to plants in control plots, leaf P concentration was 54% higher and leaf N concentration was 10% higher for plants in the P- and N-addition treatments, respectively. The N:P of leaves suggested P-limitation in the N-addition treatment and the control but not in the P-addition treatment. Root branching was highest in the P-addition treatment, and P-addition reduced mycorrhizal colonization. Conclusions: The large effect of P fertilization suggests that, like many tropical plants, P. fragrantissimum has the potential to be P-limited. Although further study is needed, we suggest that nomadic vines be added to the growth forms that respond to nutrient addition in the forest understory and conclude that nutrient-limitation seems like the rule rather than the exception in the light-limited understory

Demographic trade-offs predict tropical forest dynamics

Understanding tropical forest dynamics and planning for their sustainable management require efficient, yet accurate, predictions of the joint dynamics of hundreds of tree species. With increasing information on tropical tree life histories, our predictive understanding is no longer limited by species data but by the ability of existing models to make use of it. Using a demographic forest model, we show that the basal area and compositional changes during forest succession in a neotropical forest can be accurately predicted by representing tropical tree diversity (hundreds of species) with only five functional groups spanning two essential trade-offs—the growth-survival and stature-recruitment trade-offs. This data-driven modeling framework substantially improves our ability to predict consequences of anthropogenic impacts on tropical forests

Above- and belowground carbon stocks are decoupled in secondary tropical forests and are positively related to forest age and soil nutrients respectively

Reducing atmospheric CO2 is an international priority. One way to assist stabilising and reducing CO2 is to promote secondary tropical forest regrowth on abandoned agricultural land. However, relationships between above- and belowground carbon stocks with secondary forest age and specific soil nutrients remain unclear. Current global estimates for CO2 uptake and sequestration in secondary tropical forests focus on aboveground biomass and are parameterised using relatively coarse metrics of soil fertility. Here, we estimate total carbon stocks across a chronosequence of regenerating secondary forest stands (40–120 years old) in Panama, and assess the relationships between both above- and belowground carbon stocks with stand age and specific soil nutrients. We estimated carbon stocks in aboveground biomass, necromass, root biomass, and soil. We found that the two largest carbon pools - aboveground biomass and soil – have distinct relationships with stand age and soil fertility. Aboveground biomass contained ~61-97 Mg C ha-1 (24-39 % total carbon stocks) and significantly increased with stand age, but showed no relationship with soil nutrients. Soil carbon stocks contained ~128-206 Mg C ha-1 (52-70 % total stocks) and were unrelated to stand age, but were positively related to soil nitrogen. Root biomass carbon stocks tracked patterns exhibited by aboveground biomass. Necromass carbon stocks did not increase with stand age, but stocks were held in larger pieces of deadwood in older stands. Comparing our estimates to published data from younger and older secondary forests in the surrounding landscape, we show that soil carbon recovers within 40 years of forest regeneration, but aboveground biomass carbon stocks continue to increase past 100 years. Above- and belowground carbon stocks appear to be decoupled in secondary tropical forests. Paired measures of above- and belowground carbon stocks are necessary to reduce uncertainty in large-scale models of atmospheric CO2 uptake and storage by secondary forests

Incomplete recovery of tree community composition and rare species after 120 years of tropical forest succession in Panama

Determining how fully tropical forests regenerating on abandoned land recover characteristics of old-growth forests is increasingly important for understanding their role in conserving rare species and maintaining ecosystem services. Despite this, our understanding of forest structure and community composition recovery throughout succession is incomplete, as many tropical chronosequences do not extend beyond the first 50 years of succession. Here, we examined trajectories of forest recovery across eight 1-hectare plots in middle and later stages of forest succession (40–120 years) and five 1-hectare old-growth plots, in the Barro Colorado Nature Monument (BCNM), Panama. We first verified that forest age had a greater effect than edaphic or topographic variation on forest structure, diversity and composition and then corroborated results from smaller plots censused 20 years previously. Tree species diversity (but not species richness) and forest structure had fully recovered to old-growth levels by 40 and 90 years, respectively. However, rare species were missing, and old-growth specialists were in low abundance, in the mid- and late secondary forest plots, leading to incomplete recovery of species composition even by 120 years into succession. We also found evidence that dominance early in succession by a long-lived pioneer led to altered forest structure and delayed recovery of species diversity and composition well past a century after land abandonment. Our results illustrate the critical importance of old-growth and old secondary forests for biodiversity conservation, given that recovery of community composition may take several centuries, particularly when a long-lived pioneer dominates in early succession

A Standard Protocol for Liana Censuses

A recent increase in published studies of lianas has been paralleled by a proliferation of protocols for censusing lianas. This article seeks to increase uniformity in liana inventories by providing specific recommendations for the determination of which taxa to include, the location of diameter measurement points on individual stems, the setting of minimum stem diameter cutoffs, the treatment of multiple‐stemmed and rooted clonal groups, and the measurement of noncylindrical stems. Use of more uniform liana censusing protocols may facilitate comparison of independently collected data sets and further our understanding of global patterns in liana abundance, diversity, biomass, and dynamics

A Standard Protocol for Liana Censuses 1

A recent increase in published studies of lianas has been paralleled by a proliferation of protocols for censusing lianas. This article seeks to increase uniformity in liana inventories by providing specific recommendations for the determination of which taxa to include, the location of diameter measurement points on individual stems, the setting of minimum stem diameter cutoffs, the treatment of multiple-stemmed and rooted clonal groups, and the measurement of noncylindrical stems. Use of more uniform liana censusing protocols may facilitate comparison of independently collected data sets and further our understanding of global patterns in liana abundance, diversity, biomass, and dynamics.Peer Reviewedhttp://deepblue.lib.umich.edu/bitstream/2027.42/75009/1/j.1744-7429.2006.00134.x.pd