Landscape-level validation of allometric relationships for carbon stock estimation reveals bias driven by soil type.

Mitigation of climate change depends on accurate estimation and mapping of terrestrial carbon stocks, particularly in carbon dense tropical forests. Allometric equations can be used to robustly estimate biomass of tropical trees, but often require tree height, which is frequently unknown. Researchers and practitioners must, therefore, decide whether to directly measure a subset of tree heights to develop diameter : height (D:H) equations or rely on previously published generic equations. To date, studies comparing the two approaches have been spatially restricted and/or not randomly allocated across the landscape of interest, making the implications of deciding whether or not to measure tree heights difficult to determine. To address this issue, we use inventory data from a systematic-random forest inventory across Gabon (102 forest sites; 42,627 trees, including 7,036 height-measured trees). Using plot-specific models of D:H as a benchmark, we compare the performance of a suite of locally fitted and commonly used generic models (parameterized national, georegional, and pantropical equations) across a variety of scales, and assess which abiotic, anthropogenic, and topographical covariates contribute the most to bias in height estimation. We reveal marked spatial structure in the magnitude and direction of bias in tree height estimation using all generic models, due at least in part to soil type, which compounded to substantial error in site-level AGB estimates (of up to 38% or 150 Mg/ha). However, two generic pantropical models (Chave 2014; Feldpausch 2012) converged to within 2.5% of mean AGB at the landscape scale. Our results suggest that some (not all) pantropical equations can extrapolate AGB across large spatial scales with minimal bias in estimated mean AGB. However, extreme caution must be taken when interpreting the AGB estimates from generic models at the site-level as they fail to capture substantial spatial variation in D:H relationships, which could lead to dramatic under- or over-estimation of site-level carbon stocks. Validated allometric models derived at site- or soil-type-levels may be the best way to reduce such biases arising from landscape-level heterogeneity in D:H model fit in the Afrotropics.

Documenting success stories of management of phosphorus emissions at catchment scale: an example from the pilot river Odense, Denmark.

Documentation of the effects of different mitigation measures adopted at different scales to reduce phosphorus (P) loadings to surface waters is needed to help catchment managers select the best management practices. Water quality monitoring data from the outlets of two paired catchments (the river Odense catchment versus a neighbouring control catchment) on the island of Funen, Denmark, showed significantly different trends in annual flow-weighted P concentrations during the period 2000-2013. A significant downward trend in flow-weighted particulate P (PP) concentrations (0.051 mg P L-1) and loss (0.155 kg P ha-1) was detected for the river Odense catchment, whereas a similar trend did not emerge in the control catchment. The observed differences in PP reductions may be due to wetlands acting as P sinks since wetland restoration activities have been much more comprehensive in the river Odense catchment (1.8 ha wetlands km-2) than in the control catchment (0.5 ha wetland km-2). The excess downward trend in total P and PP in the river Odense catchment (5,600 kg P and 3,700 kg P) is corroborated by extrapolating the results from a mass-balance study and 10 years of in situ measurements of P storage (3,700 kg P and 15,000 kg P).

Successful reduction of diffuse nitrogen emissions at catchment scale: example from the pilot River Odense, Denmark.

Land-based total nitrogen (N) loadings to Danish coastal waters have been markedly reduced since 2000. This has been achieved by general measures reducing discharges from all point sources and N leaching from farmed land supplemented with more local and targeted mitigation measures such as restoration of wetlands to increase the catchment-specific N retention. In the catchment of River Odense, restoration of wetlands has been extensive. Thus, in the major gauged catchment (485 km(2)) eleven wetlands (860 ha) have been restored since 2000. A comparison of data on N concentrations and loss from a gauging station in the River Odense with data from a control catchment (772 km(2)), in which a significantly less intensive wetland restoration programme has been undertaken, showed an excess downward trend in N, amounting to 124 t N yr(-1), which can be ascribed to the intensive wetland restoration programme carried out in the River Odense catchment. In total, the N load in the River Odense has been reduced by 377 t N yr(-1) (39%) since 2000. The observed downward trend is supported by monitoring data from two wetlands restored in 2001 and 2004 in the River Odense catchment.

Vertebrate herbivory impacts seedling recruitment more than niche partitioning or density-dependent mortality.

In tropical forests, resource-based niches and density-dependent mortality are mutually compatible mechanisms that can act simultaneously to limit seedling populations. Differences in the strengths of these mechanisms will determine their roles in maintaining species coexistence. In the first assessment of these mechanisms in a Congo Basin forest, we quantified their relative strengths and tested the extent to which density-dependent mortality is driven by the distance-dependent behavior of seed and seedling predators predicted by the Janzen-Connell hypothesis. We conducted a large-scale seed addition experiment for five randomly selected tropical tree species, caging a subset of seed addition quadrats against vertebrate predators. We then developed models to assess the mechanisms that determine seedling emergence (three months after seed addition) and survival (two years after seed addition). As predicted, both niche differentiation and density-dependent mortality limited seedling recruitment, but predation had the strongest effects on seedling emergence and survival. Seedling species responded differently to naturally occurring environmental variation among sites, including variation in light levels and soil characteristics, supporting predictions of niche-based theories of tropical tree species coexistence. The addition of higher densities of seeds into quadrats initially led to greater seedling emergence, but survival to two years decreased with seed density. Seed and seedling predation reduced recruitment below levels maintained by density-dependent mortality, an indication that predators largely determine the population size of tree seedlings. Seedling recruitment was unrelated to the distance to or density of conspecific adult trees, suggesting that recruitment patterns are generated by generalist vertebrate herbivores rather than the specialized predators predicted by the Janzen-Connell hypothesis. If the role of seed and seedling predation in limiting seedling recruitment is a general phenomenon, then the relative abundances of tree species might largely depend on species-specific adaptations to avoid, survive, and recover from damage induced by vertebrate herbivores. Likewise, population declines of herbivorous vertebrate species (many of which are large and hunted) may trigger shifts in species composition of tropical forests.

Decoupling the effects of logging and hunting on an afrotropical animal community.

In tropical forests, hunting nearly always accompanies logging. The entangled nature of these disturbances complicates our ability to resolve applied questions, such as whether secondary and degraded forest can sustain populations of tropical animals. With the expansion of logging in central Africa, conservation depends on knowledge of the individual and combined impacts of logging and hunting on animal populations. Our goals were (1) to decouple the effects of selective logging and hunting on densities of animal guilds, including apes, duikers, monkeys, elephant, pigs, squirrels, and large frugivorous and insectivorous birds and (2) to compare the relative importance of these disturbances to the effects of local-scale variation in forest structure and fruit abundance. In northern Republic of Congo, we surveyed animals along 30 transects positioned in forest disturbed by logging and hunting, logging alone, and neither logging nor hunting. While sampling transects twice per month for two years, we observed 47 179 animals of 19 species and eight guilds in 1154 passages (2861 km). Species densities varied by as much as 480% among forest areas perturbed by logging and/or hunting, demonstrating the strong effects of these disturbances on populations of some species. Densities of animal guilds varied more strongly with disturbance type than with variation in forest structure, canopy cover, and fruit abundance. Independently, logging and hunting decreased density of some guilds and increased density of others: densities varied from 44% lower (pigs) to 90% higher (insectivorous birds) between logged and unlogged forest and from 61% lower (apes) to 77% higher (frugivorous birds) between hunted and unhunted forest. Their combined impacts exacerbated decreases in populations of some guilds (ape, duiker, monkey, and pig), but counteracted one another for others (squirrels, insectivorous and frugivorous birds). Together, logging and hunting shifted the relative abundance of the animal community away from large mammals toward squirrels and birds. Logged forest, even in the absence of hunting, does not maintain similar densities as unlogged forest for most animal guilds. To balance conservation with the need for economic development and wild meat in tropical countries, landscapes should be spatially managed to include protected areas, community hunting zones, and production forest.

Bushmeat supply and consumption in a tropical logging concession in northern Congo.

Unsustainable hunting of wildlife for food empties tropical forests of many species critical to forest maintenance and livelihoods of forest people. Extractive industries, including logging, can accelerate exploitation of wildlife by opening forests to hunters and creating markets for bushmeat. We monitored human demographics, bushmeat supply in markets, and household bushmeat consumption in five logging towns in the northern Republic of Congo. Over 6 years we recorded 29,570 animals in town markets and collected 48,920 household meal records. Development of industrial logging operations led to a 69% increase in the population of logging towns and a 64% increase in bushmeat supply. The immigration of workers, jobseekers, and their families altered hunting patterns and was associated with increased use of wire snares and increased diversity in the species hunted and consumed. Immigrants hunted 72% of all bushmeat, which suggests the short-term benefits of hunting accrue disproportionately to "outsiders" to the detriment of indigenous peoples who have prior, legitimate claims to wildlife resources. Our results suggest that the greatest threat of logging to biodiversity may be the permanent urbanization of frontier forests. Although enforcement of hunting laws and promotion of alternative sources of protein may help curb the pressure on wildlife, the best strategy for biodiversity conservation may be to keep saw mills and the towns that develop around them out of forests.

Logging concessions can extend the conservation estate for Central African tropical forests.

The management of tropical forest in timber concessions has been proposed as a solution to prevent further biodiversity loss. The effectiveness of this strategy will likely depend on species-specific, population-level responses to logging. We conducted a survey (749 line transects over 3450 km) in logging concessions (1.2 million ha) in the northern Republic of Congo to examine the impact of logging on large mammal populations, including endangered species such as the elephant (Loxodonta africana), gorilla (Gorilla gorilla), chimpanzee (Pan troglodytes), and bongo (Tragelaphus eurycerus). When we estimated species abundance without consideration of transect characteristics, species abundances in logged and unlogged forests were not different for most species. When we modeled the data with a hurdle model approach, however, analyzing species presence and conditional abundance separately with generalized additive models and then combining them to calculate the mean species abundance, species abundance varied strongly depending on transect characteristics. The mean species abundance was often related to the distance to unlogged forest, which suggests that intact forest serves as source habitat for several species. The mean species abundance responded nonlinearly to logging history, changing over 30 years as the forest recovered from logging. Finally the distance away from roads, natural forest clearings, and villages also determined the abundance of mammals. Our results suggest that logged forest can extend the conservation estate for many of Central Africa's most threatened species if managed appropriately. In addition to limiting hunting, logging concessions must be large, contain patches of unlogged forest, and include forest with different logging histories.

Are plant populations seed limited? A critique and meta-analysis of seed addition experiments.

We examine the relative importance of processes that underlie plant population abundance and distribution. Two opposing views dominate the field. One posits that the ability to establish at a site is determined by the availability of suitable microsites (establishment limitation), while the second asserts that recruitment is limited by the availability of seeds (seed limitation). An underlying problem is that establishment and seed limitation are typically viewed as mutually exclusive. We conducted a meta-analysis of seed addition experiments to assess the relative strength of establishment and seed limitation to seedling recruitment. We asked (1) To what degree are populations seed and establishment limited? (2) Under what conditions (e.g., habitats and life-history traits) are species more or less limited by each? (3) How can seed addition studies be better designed to enhance our understanding of plant recruitment? We found that, in keeping with previous studies, most species are seed limited. However, the effects of seed addition are typically small, and most added seeds fail to recruit to the seedling stage. As a result, establishment limitation is stronger than seed limitation. Seed limitation was greater for large-seeded species, species in disturbed microsites, and species with relatively short-lived seed banks. Most seed addition experiments cannot assess the relationship between number of seeds added and number of subsequent recruits. This shortcoming can be overcome by increasing the number and range of seed addition treatments.

Plants as reef fish: fitting the functional form of seedling recruitment.

The life histories of many species depend first on dispersal to local sites and then on establishment. After dispersal, density-independent and density-dependent mortalities modify propagule supply, determining the number of individuals that establish. Because multiple factors influence recruitment, the dichotomy of propagule versus establishment limitation is best viewed as a continuum along which the strength of propagule or establishment limitation changes with propagule input. To evaluate the relative importance of seed and establishment limitation for plants, we (1) describe the shape of the recruitment function and (2) use limitation and elasticity analyses to quantify the sensitivity of recruitment to perturbations in seed limitation and density-independent and density-dependent mortality. Using 36 seed augmentation studies for 18 species, we tested four recruitment functions against one another. Although the linear model (accounting for seed limitation and density-independent mortality) fitted the largest number of studies, the nonlinear Beverton-Holt model (accounting for density-dependent mortality) performed better at high densities of seed augmentation. For the 18 species, seed limitation constrained population size more than other sources of limitation at ambient conditions. Seedling density reached saturation with increasing seed density in many studies, but at such high densities that seedling density was primarily limited by seed availability rather than microsite availability or density dependence.

Seasonal variation in the feeding ecology of the grey-cheeked mangabey (Lophocebus albigena) in Cameroon.

Seasonal fluctuations in resource abundance often cause primates to change their feeding behavior and ecology. The objective of this study was to examine the response of a largely frugivorous monkey, the grey-cheeked mangabey (Lophocebus albigena), to seasonal variations in fruit abundance. We used 15-min scan sampling to quantify feeding, activity, and habitat use by monkeys between February and December 1998 in the Dja Reserve, Cameroon. L. albigena were found to have omnivorous feeding habits, consuming the fruits, seeds, leaves, and flowers of 132 plant species. Although monkeys fed from many plant species, only five plant species accounted for 45% of all feeding records. The number of feeding observations on a plant species was significantly correlated with its fruit production. L. albigena responded to fruit-lean periods by shifting from a diet dominated by fruit to one dominated by seeds, flowers, and young leaves. This diet shift coincided with greater use of swamp habitat and higher dietary diversity. L. albigena spent the greatest percentage of scan samples feeding and traveling, but activities varied significantly over the day. Individuals spent a significantly higher percentage of scan samples feeding during the fruit-rich season than in the fruit-lean season. Comparing our results to those of studies in Gabon and Uganda, we found that L. albigena differ across regions in the number of plant species they consume and time spent feeding. These differences may be a result of variations in tree diversity or the strength of seasonal fluctuations in resource abundance among sites.

Fruiting trees as dispersal foci in a semi-deciduous tropical forest.

Quantification of seed rain patterns is an initial step toward explaining variation in plant recruitment, and consequently, organization of forest communities. Spatially contagious patterns of seed deposition, where seeds are patchily dispersed with some sites receiving relatively high densities and others receiving low densities of seeds, may be a common phenomenon for which we have very little knowledge. For example, prior feeding events by frugivores (monkeys and birds) combined with transport and dispersal of seeds to other fruiting trees may result in the contagious deposition of non-conspecific seeds below them. Here, we examined whether fruiting trees act as dispersal foci in the semi-deciduous tropical rainforest of the Dja Reserve, Cameroon. Seed rain was sampled below the canopies of nine tree species: three typically dispersed by large, frugivorous birds, three dispersed by monkeys, and three dispersed by wind. We found no evidence that monkeys generate spatially contagious patterns of seed rain under fruiting trees at which they feed. However, we found that rates of deposition of non-conspecific seeds and species richness of seeds delivered by birds (hornbills and turacos) were significantly greater during fruiting than non-fruiting periods, and significantly greater under fruiting individuals of bird-dispersed tree species than under fruiting individuals of monkey- or wind-dispersed tree species. Additionally, during fruiting periods, the composition of non-conspecific seed rain under bird-dispersed tree species was more similar to other bird-dispersed trees than to monkey- or wind-dispersed tree species. The contagious dispersal of non-conspecific seeds to fruiting, bird-dispersed trees leads to higher seed densities under fruiting trees than those caused by local seed production. Non-conspecific seeds deposited in high densities may experience increased seed mortality even far from parent trees if predators are generalists. Alternatively, in the absence of complete density-dependent mortality, contagious seed dispersal could result in associations among species dispersed by the same dispersal agent.