Lee White

Benchmark map of forest carbon stocks in tropical regions across three continents

Developing countries are required to produce robust estimates of forest carbon stocks for successful implementation of climate change mitigation policies related to reducing emissions from deforestation and degradation (REDD). Here we present a “benchmark” map of biomass carbon stocks over 2.5 billion ha of forests on three continents, encompassing all tropical forests, for the early 2000s, which will be invaluable for REDD assessments at both project and national scales. We mapped the total carbon stock in live biomass (above- and belowground), using a combination of data from 4,079 in situ inventory plots and satellite light detection and ranging (Lidar) samples of forest structure to estimate carbon storage, plus optical and microwave imagery (1-km resolution) to extrapolate over the landscape. The total biomass carbon stock of forests in the study region is estimated to be 247 Gt C, with 193 Gt C stored aboveground and 54 Gt C stored belowground in roots. Forests in Latin America, sub-Saharan Africa, and Southeast Asia accounted for 49%, 25%, and 26% of the total stock, respectively. By analyzing the errors propagated through the estimation process, uncertainty at the pixel level (100 ha) ranged from ±6% to ±53%, but was constrained at the typical project (10,000 ha) and national (>1,000,000 ha) scales at ca. ±5% and ca. ±1%, respectively. The benchmark map illustrates regional patterns and provides methodologically comparable estimates of carbon stocks for 75 developing countries where previous assessments were either poor or incomplete.

Increasing carbon storage in intact African tropical forests

The response of terrestrial vegetation to a globally changing environment is central to predictions of future levels of atmospheric carbon dioxide. The role of tropical forests is critical because they are carbon-dense and highly productive. Inventory plots across Amazonia show that old-growth forests have increased in carbon storage over recent decades, but the response of one-third of the world’s tropical forests in Africa is largely unknown owing to an absence of spatially extensive observation networks. Here we report data from a ten-country network of long-term monitoring plots in African tropical forests. We find that across 79 plots (163 ha) above-ground carbon storage in live trees increased by 0.63 Mg C ha-1 yr-1 between 1968 and 2007 (95% confidence interval (CI), 0.22–0.94; mean interval, 1987–96). Extrapolation to unmeasured forest components (live roots, small trees, necromass) and scaling to the continent implies a total increase in carbon storage in African tropical forest trees of 0.34 Pg C yr-1 (CI, 0.15–0.43). These reported changes in carbon storage are similar to those reported for Amazonian forests per unit area, providing evidence that increasing carbon storage in old-growth forests is a pan-tropical phenomenon. Indeed, combining all standardized inventory data from this study and from tropical America and Asia together yields a comparable figure of 0.49 Mg C ha-1 yr-1 (n = 156; 562 ha; CI, 0.29–0.66; mean interval, 1987–97). This indicates a carbon sink of 1.3 Pg C yr-1 (CI, 0.8–1.6) across all tropical forests during recent decades. Taxon-specific analyses of African inventory and other data suggest that widespread changes in resource availability, such as increasing atmospheric carbon dioxide concentrations, may be the cause of the increase in carbon stocks, as some theory and models predict.

Catastrophic ape decline in western equatorial Africa

Because rapidly expanding human populations have devastated gorilla (Gorilla gorilla) and common chimpanzee (Pan troglodytes) habitats in East and West Africa, the relatively intact forests of western equatorial Africa have been viewed as the last stronghold of African apes. Gabon and the Republic of Congo alone are thought to hold roughly 80% of the worlds gorillas and most of the common chimpanzees. Here we present survey results conservatively indicating that ape populations in Gabon declined by more than half between 1983 and 2000. The primary cause of the decline in ape numbers during this period was commercial hunting, facilitated by the rapid expansion of mechanized logging. Furthermore, Ebola haemorrhagic fever is currently spreading through ape populations in Gabon and Congo and now rivals hunting as a threat to apes. Gorillas and common chimpanzees should be elevated immediately to ‘critically endangered’ status. Without aggressive investments in law enforcement, protected area management and Ebola prevention, the next decade will see our closest relatives pushed to the brink of extinction.

The primate community of the Lopé reserve, Gabon: Diets, responses to fruit scarcity, and effects on biomass

The diets of all diurnal primates (Gorilla g. gorilla, Pan t. troglodytes, Mandrillus sphinx, Colobus satanas, Cercocebus albigena, Cercopithecus nictitans, C. pogonias, C. cephus) in the Lopé Reserve, central Gabon, are described from qualitative and quantitative data collected over 10 years. A total of 397 foods were recorded, of which 91% were from plants. The diets of seven of the eight species were numerically dominated by fruit, the exception being Colobus satanas with a diet dominated by seeds. When proportion of feeding time was examined, fruit remained the dominant food class for six species, while Cercocebus albigena, like the black colobus, spent most time feeding on seeds. The number of foods recorded per species ranged from 46–220, but dietary breadth of the four species that have not been subjects of detailed study was certainly underestimated. Almost half of the foods (46%) were eaten by only one species, and only four (1%) were recorded for all eight species. At Lopé, fruit is scarce for 2–3 months during the annual dry season, and fruit crop failures can prolong this period of scarcity to as many as 8 months in some years. When fruit is scarce, the diets of all species changed to include more nonfruit foods, but chimpanzees and the three species of guenons maintained a fruit dominated diet. The total biomass of the Lopé primate community is 374 kg/km2, which is low compared to other sites in tropical Africa. Within the Lopé community, biomass correlates negatively with degree of frugivory, suggesting that fruit availability may be critical, but this does not explain the low total biomass compared to other sites. Neither floristic diversity nor the composition of the primate community at Lopé can account for the low biomass. While historical events such as climate‐induced changes in forest cover, disease, or impact of human activities cannot be ruled out, we conclude that the most likely explanation of low primate biomass is the occurrence of an ecological “catastrophe” in the fairly recent past from which populations have not yet recovered. We believe that the most likely scenario was climate‐mediated disturbance of flowering of important fruit food species. Prolonged fruit scarcity would have caused mortalities and affected the most frugivorous species more severely. Am. J. Primatol. 42:1–24, 1997.

The role of Pleistocene refugia and rivers in shaping gorilla genetic diversity in central Africa

The role of Pleistocene forest refugia and rivers in the evolutionary diversification of tropical biota has been the subject of considerable debate. A range-wide analysis of gorilla mitochondrial and nuclear variation was used to test the potential role of both refugia and rivers in shaping genetic diversity in current populations. Results reveal strong patterns of regional differentiation that are consistent with refugial hypotheses for central Africa. Four major mitochondrial haplogroups are evident with the greatest divergence between eastern (A, B) and western (C, D) gorillas. Coalescent simulations reject a model of recent east–west separation during the last glacial maximum but are consistent with a divergence time within the Pleistocene. Microsatellite data also support a similar regional pattern of population genetic structure. Signatures of demographic expansion were detected in eastern lowland (B) and Gabon/Congo (D3) mitochondrial haplogroups and are consistent with a history of postglacial expansion from formerly isolated refugia. Although most mitochondrial haplogroups are regionally defined, limited admixture is evident between neighboring haplogroups. Mantel tests reveal a significant isolation-by-distance effect among western lowland gorilla populations. However, mitochondrial genetic distances also correlate with the distance required to circumnavigate intervening rivers, indicating a possible role for rivers in partitioning gorilla genetic diversity. Comparative data are needed to evaluate the importance of both mechanisms of vicariance in other African rainforest taxa.

Nest building by lowland gorillas in the Lope Reserve, Gabon : Environmental influences and implications for censusing

We analyzed data from 373 fresh nest-sites (containing 2435 nests) of lowland gorillas (Gorilla g. gorilla)during a 4-year period in the Lopé Reserve, Gabon, to determine whether the observed variability in nest building was due to environmental influences. We recognized and defined seven types of nest in terms of the degree of construction and the raw materials used. Overall, nests built on the ground from herbaceous plants are the most common type (40%), followed by tree nests (35%). Frequencies of the different nest-types vary significantly between eight habitat-types. In habitat-types with high densities of understory herbs, ground nests predominated, but when herbs were rare, the majority of nests were in trees. A general preference for sleeping in herbaceous ground nests is indicated since trees are abundant in all habitat-types, except savanna. The frequency of nesting in trees shows a significant positive correlation with rainfall, but effects of climate are confounded by seasonal variation in use of different habitat-types. When elephants were attracted to the same localized food sources as gorillas, many tree nests were built even when herbs were available. We conclude that different nest-types reflect a variety of solutions to maximize comfort, depending on available raw materials and the probability of rainfall or disturbance by elephants or both factors. Nests are a powerful tool for population censuses and demographic studies of great apes, but problems exist in interpreting data on lowland gorilla nests. Results from this analysis show that only a third of nest-sites accurately reflects group size (of weaned individuals) and that 26% of all gorilla nest-sites could be mistaken for those of chimpanzees, as all nests, or all those visible from a transect, were in trees. Gorilla nests at Lopé were nonrandomly distributed with respect to habitat-types, and nest construction varied seasonally, thereby introducing sources of bias to transect nest counts. We discuss these problems and ones related to assessing the decay rate of nest-sites and make recommendations relevant to census work.

Why Don't Chimpanzees in Gabon Crack Nuts?

Some populations of wild chimpanzees (Pan troglodytes) use hammers and anvils of stone or wood to crack open nuts for food. Others do not. The aim of this study was to ask why one non-nut-cracking population, in the Lopé Reserve, Gabon, lacks this useful form of tool use. We tested 10 hypotheses: (1) nuts are absent; (2) nuts are few; (3) nuts are unsuitable; (4) hammers are absent; (5) hammers are unsuitable; (6) anvils are absent; (7) anvils are unsuitable; (8) nuts are displaced by better food items; (9) intelligence is insufficient; and (10) knowledge is insufficient. All but the last are clearly falsified, leaving by exclusion the likelihood that Lopés chimpanzees lack the technology—knowledge of appropriate technique—to exploit this resource. Thus, the behavioral differences across populations of these apes are cultural and not environmentally dictated. This explanation is congruent with the distribution of chimpanzee nut-cracking across Africa.

Hordes of mandrills (Mandrillus sphinx): extreme group size and seasonal male presence

Mandrill Mandrillus sphinx hordes in the LopeReserve, Gabon, the approximate centre of the mandrill species range, were studied over 3 years from 1996 to 1999. Part of the study site included gallery forests within savanna areas, allowing observation of entire hordes, hitherto impossible in dense forest habitat. Horde size and composition (sex and age classes) were documented using exact records on video film whenever a horde or subgroup crossed an open space. Mean horde size was 620, and hordes of up to 845 individuals were documented, probably the largest stable group size found in any wild, unprovisioned primate population. Hordes were cohesive throughout the study period and did not seem to be aggregations of smaller units. Mandrill societies seem to be quite different from the baboon societies, to which they have been compared to date. Mature, breeding-age males were not resident members of hordes, but entered at the onset of seasonal cycles in the females (as deduced by the presence of sexual tumescence) and emigrated once female sexual cycles ceased. The number of breeding males present in the horde at any one time is best explained by the number of sexually attractive females. It is postulated that the extreme coloration of males and strong sexual dimorphism in mandrills may have evolved through an enhanced need for competitive signals in a situation where no long-term social bonds between breeding partners exist.

Patterns of fruit-fall phenology in the Lopé Reserve, Gabon

Patterns of fruit production were monitored over a 1-year period, from June 1990–May 1991, by counting the numbers of freshly fallen ripe and unripe fruits on five 5-km line transects located within 35 km of one another, in lowland tropical rainforest in the Lope Reserve, central Gabon. A total of 195 species of fruit were found, representing at least 45 taxonomic families. The majority of fruits came from trees. Fruits were assigned to one of six categories which reflected their dispersal syndrome: succulent, arillate, dehiscent, fleshy pods, wind-dispersed, others. About three-quarters of all species had fruits characteristic of those dispersed by animals. There was a marked seasonal pattern to fruit production, with both the diversity and number of ripe fruits available peaking in January, and lowest immediately before and during the major dry season. The only climatic variable that showed a significant statistical correlation with fruit production was insolation. Candidates for the role of keystone fruit species were identified from species which fruited during the major dry season, and patterns of fruit production compared with other areas.

Mitochondrial DNA phylogeography of western lowland gorillas (Gorilla gorilla gorilla)

The geographical distribution of genetic variation within western lowland gorillas (Gorilla gorilla gorilla) was examined to clarify the population genetic structure and recent evolutionary history of this group. DNA was amplified from shed hair collected from sites across the range of the three traditionally recognized gorilla subspecies: western lowland (G. g. gorilla), eastern lowland (G. g. graueri) and mountain (G. g. beringei) gorillas. Nucleotide sequence variation was examined in the first hypervariable domain of the mitochondrial control region and was much higher in western lowland gorillas than in either of the other two subspecies. In addition to recapitulating the major evolutionary split between eastern and western lowland gorillas, phylogenetic analysis indicates a phylogeographical division within western lowland gorillas, one haplogroup comprising gorilla populations from eastern Nigeria through to southeast Cameroon and a second comprising all other western lowland gorillas. Within this second haplogroup, haplotypes appear to be partitioned geographically into three subgroups: (i) Equatorial Guinea, (ii) Central African Republic, and (iii) Gabon and adjacent Congo. There is also evidence of limited haplotype admixture in northeastern Gabon and southeast Cameroon. The phylogeographical patterns are broadly consistent with those predicted by current Pleistocene refuge hypotheses for the region and suggest that historical events have played an important role in shaping the population structure of this subspecies.