Browsing by Author "Clark, CJ"

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Open Access
Decoupling the effects of logging and hunting on an afrotropical animal community.
(Ecol Appl, 2011-07) Poulsen, JR; Clark, CJ; Bolker, BM
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.
Open Access
Diet selection is related to breeding status in two frugivorous hornbill species of Central Africa
(Journal of Tropical Ecology, 2014-07) Lamperti, AM; French, AR; Dierenfeld, ES; Fogiel, MK; Whitney, KD; Stauffer, DJ; Holbrook, KM; Hardesty, BD; Clark, CJ; Poulsen, JR; Wang, BC; Smith, TB; Parker, VT
Avian diet selection is hypothesized to be sensitive to seasonal changes in breeding status, but few tests exist for frugivorous tropical birds. Frugivorous birds provide an interesting test case because fruits are relatively deficient in minerals critical for reproduction. Here, we quantify annual patterns of fruit availability and diet for two frugivorous hornbill (Bucerotidae) species over a 5.5-y period to test for patterns of diet selection. Data from the lowland tropical rain forest of the Dja Reserve, Cameroon, are used to generate two nutritional indices. One index estimates the nutrient concentration of the diet chosen by Ceratogymna atrata and Bycanistes albotibialis on a monthly basis using 3165 feeding observations combined with fruit pulp sample data. The second index is an estimate of nutrient concentration of a non-selective or neutral diet across the study area based on tree fruiting phenology, vegetation survey and fruit-pulp sample data. Fifty-nine fruit pulp samples representing 40 species were analysed for 16 nutrient categories to contribute to both indices. Pulp samples accounted for approximately 75% of the observed diets. The results support expected patterns of nutrient selection. The two hornbill species selected a diet rich in calcium during the early breeding season (significantly so for B. albotibialis in July and August). Through the brooding and fledging periods, they switched from a calcium-rich diet to one rich in iron and caloric content as well as supplemental protein in the form of invertebrates. Calcium, the calcium to phosphorus ratio and fat concentration were the strongest predictors of breeding success (significant for calcium and Ca:P for B. albotibialis in June). We conclude that hornbills actively select fruit based on nutritional concentration and mineral concentration and that the indices developed here are useful for assessing frugivore diet over time. Copyright © Cambridge University Press 2014.
Open Access
From town to national park: Understanding the long-term effects of hunting and logging on tree communities in Central Africa
(Forest Ecology and Management, 2021-11-01) Maicher, V; Clark, CJ; Harris, DJ; Medjibe, VP; Poulsen, JR
Anthropogenic disturbances are changing the structure and composition of tropical forests worldwide. Multiple disturbances often occur simultaneously in forests. Hunting and logging, for example, are within-forest disturbances that impact vast areas of seemingly intact rainforests. Despite recent work on the individual effects of these disturbances, our understanding of how they interact to influence tree communities is still limited. In northern Republic of Congo, we explored the effects of hunting and logging on tree communities. Over an 8-year period, we monitored 12,552 tree stems (≥10 cm diameter-at-breast height) spread over 30 1-ha plots along a gradient of human disturbance to compare the tree diversity between hunted and logged forest, once-logged forest, and protected forest free of both disturbances. Tree density, species richness, and community composition were affected by both hunting and logging. Forest close to human settlements was richer, more heterogenous, and more dynamic in species composition across censuses. In hunted and logged forest, fast-growing secondary species with low shade tolerance replaced old growth species. Comparatively, the once-logged forest had the greatest stem density and intermediate species richness with an increased density of shade-bearing species over time. Both tree species spatial turnover and tree recruitment were greatly affected by proximity to human settlements. A shift towards abiotically dispersed trees and increasing seed predation by rodents near villages can partly explain the differences in tree recruitment across the forest types. The combination of hunting and logging seems to have a greater impact on tree communities than either single disturbance, especially with nearness to villages.
Open Access
Long Distance Seed Dispersal by Forest Elephants
(Frontiers in Ecology and Evolution, 2021-12-22) Poulsen, JR; Beirne, C; Rundel, C; Baldino, M; Kim, S; Knorr, J; Minich, T; Jin, L; Núñez, CL; Xiao, S; Mbamy, W; Obiang, GN; Masseloux, J; Nkoghe, T; Ebanega, MO; Clark, CJ; Fay, MJ; Morkel, P; Okouyi, J; White, LJT; Wright, JP
By dispersing seeds long distances, large, fruit-eating animals influence plant population spread and community dynamics. After fruit consumption, animal gut passage time and movement determine seed dispersal patterns and distances. These, in turn, are influenced by extrinsic, environmental variables and intrinsic, individual-level variables. We simulated seed dispersal by forest elephants (Loxodonta cyclotis) by integrating gut passage data from wild elephants with movement data from 96 individuals. On average, elephants dispersed seeds 5.3 km, with 89% of seeds dispersed farther than 1 km. The longest simulated seed dispersal distance was 101 km, with an average maximum dispersal distance of 40.1 km. Seed dispersal distances varied among national parks, perhaps due to unmeasured environmental differences such as habitat heterogeneity and configuration, but not with human disturbance or habitat openness. On average, male elephants dispersed seeds farther than females. Elephant behavioral traits strongly influenced dispersal distances, with bold, exploratory elephants dispersing seeds 1.1 km farther than shy, idler elephants. Protection of forest elephants, particularly males and highly mobile, exploratory individuals, is critical to maintaining long distance seed dispersal services that shape plant communities and tropical forest habitat.
Open Access
Natural regeneration of selected timber species in the Republic of Congo
(African Journal of Ecology, 2014) Medjibe, VP; Poulsen, JR; Clark, CJ; Mbani, OA
© 2014 John Wiley & Sons Ltd.Natural regeneration of timber species is critical to the sustainable management of tropical forests. To understand what determines regeneration success of timber species in the Congo Basin, we evaluated whether seedling recruitment rates differed between forest logged 30 years previously and unlogged forest and determined the environmental factors that influence seedling density, growth and survival. We monitored the fate of 2186 seedlings of seven timber species within 462, 25-m<sup>2</sup> plots located along 21 transects. We characterized seedling plots by light availability, soil nutrient availability and pH, and abundance of mammalian herbivores and then used linear and generalized linear mixed models to evaluate the variables that influenced seedling density, growth and survival. Light availability and canopy openness were 18% and 81% higher in logged than unlogged forest, and concentration of soil nutrients varied between sites. Seedling density was 32% higher in unlogged than logged forest. Taking all species together, seedling survival was positively correlated with calcium and negatively with magnesium and available phosphorus. Rates of seedling growth increased with available light. Taken separately, seedlings of the selected timber species responded differently to abiotic and biotic factors, demonstrating species-specific regeneration requirements.
Open Access
Old growth Afrotropical forests critical for maintaining forest carbon
(Global Ecology and Biogeography, 2020-10-01) Poulsen, JR; Medjibe, VP; White, LJT; Miao, Z; Banak-Ngok, L; Beirne, C; Clark, CJ; Cuni-Sanchez, A; Disney, M; Doucet, JL; Lee, ME; Lewis, SL; Mitchard, E; Nuñez, CL; Reitsma, J; Saatchi, S; Scott, CT
Aim: Large trees [≥ 70 cm diameter at breast height (DBH)] contribute disproportionately to aboveground carbon stock (AGC) across the tropics but may be vulnerable to changing climate and human activities. Here we determine the distribution, drivers and threats to large trees and high carbon forest. Location: Central Africa. Time period: Current. Major taxa studied: Trees. Methods: Using Gabon's new National Resource Inventory of 104 field sites, AGC was calculated from 67,466 trees from 578 species and 97 genera. Power and Michaelis–Menten models assessed the contribution of large trees to AGC. Environmental and anthropogenic drivers of AGC, large trees, and stand variables were modelled using Akaike’s information criterion (AIC) weights to calculate average regression coefficients for all p. ossible models. Results: Mean AGC for trees ≥ 10 cm DBH in Gabonese forestlands was 141.7 Mg C/ha, with averages of 166.6, 171.3 and 96.6 Mg C/ha in old growth, concession and secondary forest. High carbon forests occurred where large trees are most abundant: 31% of AGC was stored in large trees (2.3% of all stems). Human activities largely drove variation in AGC and large trees, but climate and edaphic conditions also determined stand variables (basal area, tree height, wood density, stem density). AGC and large trees increased with distance from human settlements; AGC was 40% lower in secondary than primary and concession forests and 33% higher in protected than non-managed areas. Main conclusions: AGC and large trees were negatively associated with human activities, highlighting the importance of forest management. Redefining large trees as ≥ 50 cm DBH (4.3% more stems) would account for 20% more AGC. This study demonstrates that protecting relatively undisturbed forests can be disproportionately effective in conserving carbon and suggests that including sustainable forestry in programs like reduced emissions for deforestation and forest degradation could maintain carbon dense forests in logging concessions that are a large proportion of remaining Central African forests.
Open Access
Plants as reef fish: fitting the functional form of seedling recruitment.
(Am Nat, 2007-08) Poulsen, JR; Osenberg, CW; Clark, CJ; Levey, DJ; Bolker, BM
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.
Open Access
Seed dispersal by a diurnal primate community in the Dja Reserve, Cameroon
(Journal of Tropical Ecology, 2001-11) Poulsen, JR; Clark, CJ; Smith, TB
Open Access
Using Model Analysis to Unveil Hidden Patterns in Tropical Forest Structures
(Frontiers in Ecology and Evolution, 2021-11-30) Picard, N; Mortier, F; Ploton, P; Liang, J; Derroire, G; Bastin, JF; Ayyappan, N; Bénédet, F; Boyemba Bosela, F; Clark, CJ; Crowther, TW; Engone Obiang, NL; Forni, É; Harris, D; Ngomanda, A; Poulsen, JR; Sonké, B; Couteron, P; Gourlet-Fleury, S
When ordinating plots of tropical rain forests using stand-level structural attributes such as biomass, basal area and the number of trees in different size classes, two patterns often emerge: a gradient from poorly to highly stocked plots and high positive correlations between biomass, basal area and the number of large trees. These patterns are inherited from the demographics (growth, mortality and recruitment) and size allometry of trees and tend to obscure other patterns, such as site differences among plots, that would be more informative for inferring ecological processes. Using data from 133 rain forest plots at nine sites for which site differences are known, we aimed to filter out these patterns in forest structural attributes to unveil a hidden pattern. Using a null model framework, we generated the anticipated pattern inherited from individual allometric patterns. We then evaluated deviations between the data (observations) and predictions of the null model. Ordination of the deviations revealed site differences that were not evident in the ordination of observations. These sites differences could be related to different histories of large-scale forest disturbance. By filtering out patterns inherited from individuals, our model analysis provides more information on ecological processes.
Open Access
Vertebrate herbivory impacts seedling recruitment more than niche partitioning or density-dependent mortality.
(Ecology, 2012-03) Clark, CJ; Poulsen, JR; Levey, DJ
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.

Browsing by Author "Clark, CJ"

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