Changes in tree community structure in defaunated forests are not driven only by dispersal limitation

1. Bushmeat hunting has reduced population sizes of large frugivorous vertebrates throughout the tropics, thereby reducing the dispersal of seeds. This is believed to affect tree population dynamics, and therefore community composition, because the seed dispersal of large‐seeded trees depends upon large‐bodied vertebrates.
2. We report on a long‐running study of the effect of defaunation on a tropical tree community. In three censuses over 11 years, we compared sapling recruitment between a hunted and a nonhunted site, which are nearby and comparable to one another, to determine the extent to which species composition has changed through time following defaunation. We expected to find a reduced abundance of tree species that rely on large frugivores for dispersal at the hunted site and altered community structure as a consequence.
3. Although community composition at the hunted site diverged from that at the nonhunted site, the changes were independent of dispersal syndrome, with no trend toward a decline in species that are dispersed by large, hunted vertebrates. Moreover, the loss of large‐bodied dispersers did not generate the changes in tree community composition that we hypothesized. Some species presumed to rely on large‐bodied frugivores for dispersal are effectively recruiting despite the absence of their dispersers.
4. Synthesis: The presumption that forests depleted of large‐bodied dispersers will experience rapid, directional compositional change is not fully supported by our results. Altered species composition in the sapling layer at the hunted site, however, indicates that defaunation may be connected with changes to the tree community, but that the nature of these changes is not unidirectional as previously assumed. It remains difficult to predict how defaunation will affect tree community composition without a deeper understanding of the driving mechanisms at play.

     

Bushmeat hunting changes regeneration of African rainforests

To assess ecological consequences of bushmeat hunting in African lowland rainforests, we compared paired sites, with high and low hunting pressure, in three areas of southeastern Nigeria. In hunted sites, populations of important seed dispersers—both small and large primates (including the Cross River gorilla, Gorilla gorilla diehli)—were drastically reduced. Large rodents were more abundant in hunted sites, even though they are hunted. Hunted and protected sites had similar mature tree communities dominated by primate-dispersed species. In protected sites, seedling communities were similar in composition to the mature trees, but in hunted sites species with other dispersal modes dominated among seedlings. Seedlings emerging 1 year after clearing of all vegetation in experimental plots showed a similar pattern to the standing seedlings. This study thus verifies the transforming effects of bushmeat hunting on plant communities of tropical forests and is one of the first studies to do so for the African continent.     

Bushmeat and the Fate of Trees with Seeds Dispersed by Large Primates in a Lowland Rain Forest in Western Amazonia

In Neotropical forests, large fruit-eating primates play important ecological roles as dispersal agents of large seeds. Bushmeat hunting threatens to disrupt populations of primates and large-seeded trees. We test the hypothesis that otherwise intact Neotropical forests with depressed populations of large primates experience decline in recruitment of large-seeded trees. We quantify the proportion of small juveniles (> 0.5 m tall–1 cm diameter at breast height, DBH) of large primate-dispersed tree species found underneath heterospecifc trees that are also dispersed by large primates at two protected sites in Manu National Park and one hunted site outside Manu N.P. in southeastern Peru. The forests are comparable in edaphic and climatic qualities, successional stage, and adult tree species composition. We found that hunting locally exterminates populations of large primates, and reduced primates of intermediate body size (hereafter "medium primates") by 80 percent. Moreover, tree species richness was 55 percent lower and density of species dispersed by large and medium-bodied primates 60 percent lower in hunted than in protected sites. In addition, richness and density of abiotically dispersed species and plants dispersed by non-game animals are greater in hunted sites. Overhunting threatens to disrupt the ecological interactions between primates and the plants that rely on them for seed dispersal and recruitment. Sustainable wildlife management plans are urgently needed, because protected areas are at risk of becoming "island" parks if buffer zones become empty of animals and have impoverished flora.     

The Plight of Large Animals in Tropical Forests and the Consequences for Plant Regeneration

We introduce a special section that addresses the bushmeat or wild meat crisis, its direct impact on game species, and its indirect impact on plants in tropical forests.     

Experimental defaunation of terrestrial mammalian herbivores alters tropical rainforest understorey diversity

It has been suggested that tropical defaunation may unleash community-wide cascading effects, leading to reductions in plant diversity. However, experimental evidence establishing cause–effect relationships thereof is poor. Through a 5 year exclosure experiment, we tested the hypothesis that mammalian defaunation affects tree seedling/sapling community dynamics leading to reductions in understorey plant diversity. We established plot triplets (n = 25) representing three defaunation contexts: terrestrial-mammal exclosure (TE), medium/large mammal exclosure (PE) and open access controls (C). Seedlings/saplings 30–100 cm tall were marked and identified within each of these plots and re-censused three times to record survival and recruitment. In the periods 2010–2011 and 2011–2013, survival was greater in PE than in C plots and recruitment was higher in TE plots than in C plots. Overall, seedling density increased by 61% in TE plots and 23% in PE plots, whereas it decreased by 5% in C plots. Common species highly consumed by mammals (e.g. Brosimum alicastrum and Ampelocera hottlei) increased in their abundance in TE plots. Rarefaction curves showed that species diversity decreased in TE plots from 2008 to 2013, whereas it remained similar for C plots. Given the prevalence of tropical defaunation, we posit this is an anthropogenic effect threatening the maintenance of tropical forest diversity.     

Loss of animal seed dispersal increases extinction risk in a tropical tree species due to pervasive negative density dependence across life stages

Overhunting in tropical forests reduces populations of vertebrate seed dispersers. If reduced seed dispersal has a negative impact on tree population viability, overhunting could lead to altered forest structure and dynamics, including decreased biodiversity. However, empirical data showing decreased animal-dispersed tree abundance in overhunted forests contradict demographic models which predict minimal sensitivity of tree population growth rate to early life stages. One resolution to this discrepancy is that seed dispersal determines spatial aggregation, which could have demographic consequences for all life stages. We tested the impact of dispersal loss on population viability of a tropical tree species, Miliusa horsfieldii, currently dispersed by an intact community of large mammals in a Thai forest. We evaluated the effect of spatial aggregation for all tree life stages, from seeds to adult trees, and constructed simulation models to compare population viability with and without animal-mediated seed dispersal. In simulated populations, disperser loss increased spatial aggregation by fourfold, leading to increased negative density dependence across the life cycle and a 10-fold increase in the probability of extinction. Given that the majority of tree species in tropical forests are animal-dispersed, overhunting will potentially result in forests that are fundamentally different from those existing now.     

Bushmeat hunting in the Congo Basin: an assessment of impacts and options for mitigation

Hunting of wild animals is an important component of household economies in the Congo Basin. Results from the growing corpus of quantitative studies show that: a) bushmeat remains the primary source of animal protein for the majority of Congo Basin families; b) bushmeat hunting can constitute a significant source of revenue for forest families; c) bushmeat consumption by low density populations living in the forest may be sustainable at present; d) demand for bushmeat by growing numbers of urban consumers has created a substantial market for bushmeat that is resulting in a halo of defaunation around population centres, and may be driving unsustainable levels of hunting, even in relatively isolated regions; and e) large bodied animals with low reproductive rates are most susceptible to over-exploitation compared with more r-selected species that apparently can tolerate relatively intensive hunting (Mangel et al. 1996). As urban populations continue to grow and economies revitalise, unless action is taken to alter the demand for, and the supply of bushmeat, the forests of the Congo Basin will be progressively stripped of certain wildlife species, risking their extirpation or extinction, and the loss of values they confer to local economies. Consequently, it is essential that a) logging companies are encouraged or coerced not to facilitate bushmeat hunting and transportation in their concessions, b) we develop a better understanding of the elasticity of bushmeat demand, c) that pilot bushmeat substitution projects are supported and their impact on demand evaluated, and d) social marketing activities are put in place to attempt to direct consumer preferences for animal protein away from bushmeat species that are particularly susceptible to over-exploitation.     

The Role of Mammals in Creating and Modifying Seedshadows in Tropical Forests and Some Possible Consequences of Their Elimination

Mammal populations are increasingly hunted, yet the consequences of their disappearance from tropical forests have only recently been explored. Here, we summarize current research on the role of mammals in seed dispersal and postdispersal processes, such as seed predation and secondary dispersal, in different tropical regions. We evaluate how mammal features influence seedshadows and ultimately forest regeneration. Finally, we discuss the potential effect of changes in seedshadows caused by the elimination of many medium- and large-sized mammals. The complex role that mammals play in creating and modifying seedshadows in tropical forests cannot be easily quantified, and in this review we emphasize the variation that exists both within and among mammal taxa and across continents. To bridge this gap in information, we suggest that more studies should evaluate the relative importance of the disappearance of both seed dispersers and seed predators for particular plant species so that we may begin to understand the balance between these two influences. We also suggest that future studies identify ecological redundancy in nonhunted vertebrates within any particular community to evaluate compensatory behavior that may help ameliorate some of the negative effects of hunting of large and medium mammals.     

Rainforest seed rain into abandoned tropical Australian pasture is dependent on adjacent rainforest structure and extent

It is well known that the recovery of abandoned tropical pastures to secondary rainforest benefits from the arrival of seeds from adjacent rainforest patches. Less is known, however, about how the structural attributes of adjacent rainforest (e.g. tree density, canopy cover and tree height) impact seed rain patterns into abandoned pastures. Between 2011 and 2013, we used seed traps and ground seed surveys to track the richness and abundance of rainforest seeds entering abandoned pastures in Australia's wet tropics. We also tested how seed rain diversity is related to the distance from forest, the proportion of forest cover in the landscape and several structural attributes of adjacent forest patches, specifically average tree height, canopy cover, tree species richness and density. Almost no seeds were captured in elevated pasture seed traps, even near forest remnants. Abundant forest seeds were found in ground surveys but only within 10 m of forest edges. In ground surveys, seeds from wind‐dispersed species were more abundant, but less species rich, than animal‐dispersed species. A survey of pasture seedling recruits suggested that some forest seeds must be dispersing more than 10 m into pasture at very low frequencies, but only a few species are establishing there. Recruits were predominantly animal‐dispersed not wind‐dispersed species. In addition to distance from forest and the proportion of forest within a 100‐ to 200‐m radius of sampling sites, the richness and density of adjacent forest trees were the most important factors for explaining the probability of seed occurrence in abandoned pastures. Results suggest that without some restoration assistance, the recovery of abandoned pastures into secondary rainforest in Australia's tropical rainforests will likely be limited, at least in part, by a very low rate of seed dispersal away from forest edges and by the diversity and density of trees in adjacent remnant forests.     

Pattern and sustainability of the bushmeat trade in the Haut Niger National Park,Republic of Guinea

To document the bushmeat trade in the buffer zone of the Haut Niger National Park (HNNP), Republic of Guinea, we monitored the number of carcasses offered for sale in 16 villages over a one year period in 2001. Overall, we counted a total of 9134 carcasses from 27 species. This represents a biomass harvest of about 170,000 kg, a figure that could reach 335,100 kg when game consumed by hunters (i.e. not sold) and markets not sampled are taken into account. Ungulates (eleven species) and rodents (four species) accounted for 53.5% and 24.8% of the carcasses counted respectively (80.8% and 11.4% of the total biomass, respectively). The number of carcasses counted in villages bordering a heavily trafficked road was significantly higher than in more rural villages, thus highlighting the stimulating effect of roads on the bushmeat trade. Compared with existing data collected in 1995, the proportion of ungulates counted in the Mansira Moribaya village in 2001 increased significantly while that of rodents decreased significantly. The harvest of duiker species at Mansira Moribaya was in excess of the maximum sustainable harvest. These results of this study were used to establish a sustainable wildlife management scheme in the HNNP.     

Population history and gene dispersal inferred from spatial genetic structure of a Central African timber tree,Distemonanthus benthamianus (Caesalpinioideae)

African rainforests have undergone major distribution range shifts during the Quaternary, but few studies have investigated their impact on the genetic diversity of plant species and we lack knowledge on the extent of gene flow to predict how plant species can cope with such environmental changes. Analysis of the spatial genetic structure (SGS) of a species is an effective method to determine major directions of the demographic history of its populations and to estimate the extent of gene dispersal. This study characterises the SGS of an African tropical timber tree species, Distemonanthus benthamianus, at various spatial scales in Cameroon and Gabon. Displaying a large continuous distribution in the Lower Guinea domain, this is a model species to detect signs of past population fragmentation and recolonization, and to estimate the extent of gene dispersal. Ten microsatellite loci were used to genotype 295 adult trees sampled from eight populations. Three clearly differentiated gene pools were resolved at this regional scale and could be linked to the biogeographical history of the region, rather than to physical barriers to gene flow. A comparison with the distribution of gene pools observed for two other tree species living in the same region invalidates the basic assumption that all species share the same Quaternary refuges and recolonization pathways. In four populations, significant and similar patterns of SGS were detected. Indirect estimates of gene dispersal distances (sigma) obtained for three populations ranged from 400 to 1200 m, whereas neighbourhood size estimates ranged from 50 to 110.     

五指山热带雨林黑桫椤种群及其所在群落特征

黑桫椤(Alsophila podophylla)属国家二级保护植物.作者在海南岛五指山山地雨林和低地雨林分别设立样地对黑桫椤种群及其所在群落进行了调查,分析了热带雨林黑桫椤种群的年龄结构、空间分布格局、数量分布以及群落结构和物种多样性.结果表明,山地雨林和低地雨林中黑桫椤种群分别为稳定型种群和增长型种群,都是完全集群分布.低地雨林的黑桫椤平均密度是山地雨林的1.7倍.土壤pH值对黑桫椤种群密度有负的影响,全磷含量对其有正的影响.山地雨林林地土壤pH值大而全磷含量低,低地雨林刚好相反.土壤酸碱度和全磷含量的差异是导致两地黑桫椤数量差异的原因之一.pH值对黑桫椤平均密度的影响效应是全磷含量的2.5倍,说明pH值是影响黑桫椤种群密度的主要因素.黑桫椤所在群落垂直结构分为3个乔木亚层、2个灌木亚层及草本层.山地雨林黑桫椤所在群落的Simpson指数和Shannon-Wiener指数分别为0.957和5.305,均匀度为0.811;低地雨林黑桫椤所在群落的Simpson指数和Shannon-Wiener指数分别为0.937和5.484,均匀度为0.804.总之,五指山山地雨林和低地雨林的黑桫椤种群年龄结构、分布格局和数量分布都有一定差异,这些差异是由生境差异造成的.热带雨林黑桫椤所在群落结构复杂,物种多样性丰富.     

Fine-scale genetic structure and gene dispersal inferences in 10 neotropical tree species

The extent of gene dispersal is a fundamental factor of the population and evolutionary dynamics of tropical tree species, but directly monitoring seed and pollen movement is a difficult task. However, indirect estimates of historical gene dispersal can be obtained from the fine-scale spatial genetic structure of populations at drift-dispersal equilibrium. Using an approach that is based on the slope of the regression of pairwise kinship coefficients on spatial distance and estimates of the effective population density, we compare indirect gene dispersal estimates of sympatric populations of 10 tropical tree species. We re-analysed 26 data sets consisting of mapped allozyme, SSR (simple sequence repeat), RAPD (random amplified polymorphic DNA) or AFLP (amplified fragment length polymorphism) genotypes from two rainforest sites in French Guiana. Gene dispersal estimates were obtained for at least one marker in each species, although the estimation procedure failed under insufficient marker polymorphism, limited sample size, or inappropriate sampling area. Estimates generally suffered low precision and were affected by assumptions regarding the effective population density. Averaging estimates over data sets, the extent of gene dispersal ranged from 150 m to 1200 m according to species. Smaller gene dispersal estimates were obtained in species with heavy diaspores, which are presumably not well dispersed, and in populations with high local adult density. We suggest that limited seed dispersal could indirectly limit effective pollen dispersal by creating higher local tree densities, thereby increasing the positive correlation between pollen and seed dispersal distances. We discuss the potential and limitations of our indirect estimation procedure and suggest guidelines for future studies.     

海南岛甘什岭热带低地雨林棕榈藤空间点格局分析

以海南岛甘什岭热带低地雨林白藤(Calamus tetradactylus)、大白藤(C.faberii)、华南省藤(C.rhabdocladus)、黄藤(Daemonorops margaritae)和小钩叶藤(Plectocomia microstachys)5种棕榈藤种群为研究对象,采用点格局分析方法对棕榈藤种内和种间空间分布格局及相互关系进行分析。结果表明:(1)海南岛甘什岭热带低地雨林棕榈藤种群的空间格局与空间尺度有着密切的关系,在小尺度上,5种棕榈藤种群都更倾向于聚集分布,随着尺度的增长,各棕榈藤种群主要趋向于随机分布,形成不同的空间分布格局。(2)各藤种间的空间关联情况在整体尺度上表现出明显负关联的是白藤和黄藤、白藤和大白藤,这可能是因为它们对生存空间及养分的需求相同或相似,存在明显的竞争关系。(3)华南省藤和大白藤在整体尺度上表现出明显正关联,这可能是由于它们在生长过程中相互庇护,用于抵御动物捕食和热带雨林内多变的外部环境。(4)黄藤和小钩叶藤以及白藤和小钩叶藤呈现出明显不相关,这说明它们对环境和生存空间的依赖没有冲突性,能做到和平共存。     

Characterization of animal communities involved in seed dispersal and predation of Guibourtia tessmannii (Harms) J.Léonard,a species newly listed on Appendix II of CITES

Characterization of the ecology of endangered timber species is a crucial step in any forest management strategy. In this study, we described the animal communities involved in seed dispersal and predation of a high‐value timber species Guibourtia tessmannii (Fabaceae; Detarioideae), which is newly listed on Appendix II of CITES. We compared the animal communities between two forest sites (Bambidie in Gabon and Ma'an in Cameroon). A total of 101 hr of direct observations and 355 days of camera trapping revealed that a primate (Cercopithecus nictitans nictitans) and a hornbill (Ceratogymna atrata) were important seed dispersers in Gabon. Conversely, a greater presence of a rodent (Cricetomys emini), which could act both as predator and disperser, was observed in Cameroon. This study suggests that animal communities involved in seed dispersal of G. tessmannii may vary depending on environmental conditions and anthropogenic impacts. However, further studies are needed to properly identify the factors involved in seed dispersal and predation of G. tessmannii.     

Low levels of realized seed and pollen gene flow and strong spatial genetic structure in a small,isolated and fragmented population of the tropical tree Copaifera langsdorffii Desf

Over the past century, the Brazilian Atlantic forest has been reduced to small, isolated fragments of forest. Reproductive isolation theories predict a loss of genetic diversity and increases in inbreeding and spatial genetic structure (SGS) in such populations. We analysed eight microsatellite loci to investigate the pollen and seed dispersal patterns, genetic diversity, inbreeding and SGS of the tropical tree Copaifera langsdorffii in a small (4.8 ha), isolated population. All 112 adult trees and 128 seedlings found in the stand were sampled, mapped and genotyped. Seedlings had significantly lower levels of genetic diversity (A=16.5±0.45, mean±95% s.e.; H_e=0.838±0.006) than did adult trees (A=23.2±0.81; H_e=0.893±0.030). Parentage analysis did not indicate any seed immigration (m_seeds=0) and the pollen immigration rate was very low (m_pollen=0.047). The average distance of realized pollen dispersal within the stand was 94 m, with 81% of the pollen travelling <150 m. A significant negative correlation was found between the frequency and distance of pollen dispersal (r=−0.79, P<0.01), indicating that short-distance pollinations were more frequent. A significant SGS for both adults (∼50 m) and seedlings (∼20 m) was also found, indicating that most of the seeds were dispersed over short distances. The results suggested that the spatial isolation of populations by habitat fragmentation can restrict seed and pollen gene flow, increase SGS and affect the genetic diversity of future generations.     

Predicting the Long-Term Effects of Hunting on Plant Species Composition and Diversity in Tropical Forests

Hunting can change abundances of vertebrate seed predators and seed dispersers, causing species‐specific changes in seed dispersal and seed predation and altering seedling communities. What are the consequences of these changes for the adult plant community in the next generation and beyond? Here, I derive equations showing how reduced seed dispersal reduces plant reproduction by intensifying kin competition, increasing vulnerability to natural enemies, and reducing the proportion of seeds passing through disperser guts. I parameterize these equations with available empirical data to estimate the likely effects on next‐generation abundances. I then consider the indirect effects and longer‐term feedbacks of changed seed or adult abundances on reproductive rates due to density‐dependent interactions with natural enemies and mutualists, as well as niche differentiation with competitors, and discuss their likely qualitative effects. The factors limiting seed disperser and seed predator populations in natural and hunted forests emerge as critical for determining the long‐term effects of hunting on rates of seed dispersal and seed predation. For example, where seed dispersers are held to a constant abundance by hunters, decreases in the availability of their preferred food plants are expected to lead to increased per‐seed dispersal probabilities, potentially to the point of compensating for the initial disperser decline. I close by discussing the likely reversibility of hunting‐induced changes in tropical forests and key questions and directions for future research.