Shifting mammal communities and declining species richness along an elevational gradient on Mount Kenya.

Conservation areas encompassing elevation gradients are biodiversity hotspots because they contain a wide range of habitat types in a relatively small space. Studies of biodiversity patterns along elevation gradients, mostly on small mammal or bird species, have documented a peak in diversity at mid elevations. Here, we report on a field study of medium and large mammals to examine the impact of elevation, habitat type, and gross primary productivity on community structure. Species richness was observed using a camera trap transect with 219 sites situated across different habitat types from 2329 to 4657 m above the sea level on the western slope of Mt Kenya, the second highest mountain in Africa. We found that the lowest elevation natural habitats had the highest species richness and relative abundance and that both metrics decreased steadily as elevation increased, paralleling changes in gross primary productivity, and supporting the energy richness hypothesis. We found no evidence for the mid-domain effect on species diversity. The lowest elevation degraded Agro-Forestry lands adjacent to the National Park had high activity of domestic animals and reduced diversity and abundance of native species. The biggest difference in community structure was between protected and unprotected areas, followed by more subtle stepwise differences between habitats at different elevations. Large carnivore species remained relatively consistent but dominant herbivore species shifted along the elevation gradient. There was some habitat specialization and turnover in species, such that the elevation gradient predicts a high diversity of species, demonstrating the high conservation return for protecting mountain ecosystems for biodiversity conservation.

A unified explanation for the morphology of raised peatlands.

Raised peatlands, or bogs, are gently mounded landforms that are composed entirely of organic matter1-4 and store the most carbon per area of any terrestrial ecosystem5. The shapes of bogs are critically important because their domed morphology4,6,7 accounts for much of the carbon that bogs store and determines how they will respond to interventions8,9 to stop greenhouse gas emissions and fires after anthropogenic drainage10-13. However, a general theory to infer the morphology of bogs is still lacking4,6,7. Here we show that an equation based on the processes universal to bogs explains their morphology across biomes, from Alaska, through the tropics, to New Zealand. In contrast to earlier models of bog morphology that attempted to describe only long-term equilibrium shapes4,6,7 and were, therefore, inapplicable to most bogs14-16, our approach makes no such assumption and makes it possible to infer full shapes of bogs from a sample of elevations, such as a single elevation transect. Our findings provide a foundation for quantitative inference about the morphology, hydrology and carbon storage of bogs through Earth's history, as well as a basis for planning natural climate solutions by rewetting damaged bogs around the world.

Holocene bidirectional river system along the Kenya Rift and its influence on East African faunal exchange and diversity gradients.

East Africa is a global biodiversity hotspot and exhibits distinct longitudinal diversity gradients from west to east in freshwater fishes and forest mammals. The assembly of this exceptional biodiversity and the drivers behind diversity gradients remain poorly understood, with diversification often studied at local scales and less attention paid to biotic exchange between Afrotropical regions. Here, we reconstruct a river system that existed for several millennia along the now semiarid Kenya Rift Valley during the humid early Holocene and show how this river system influenced postglacial dispersal of fishes and mammals due to its dual role as a dispersal corridor and barrier. Using geomorphological, geochronological, isotopic, and fossil analyses and a synthesis of radiocarbon dates, we find that the overflow of Kenyan rift lakes between 12 and 8 ka before present formed a bidirectional river system consisting of a "Northern River" connected to the Nile Basin and a "Southern River," a closed basin. The drainage divide between these rivers represented the only viable terrestrial dispersal corridor across the rift. The degree and duration of past hydrological connectivity between adjacent river basins determined spatial diversity gradients for East African fishes. Our reconstruction explains the isolated distribution of Nilotic fish species in modern Kenyan rift lakes, Guineo-Congolian mammal species in forests east of the Kenya Rift, and recent incipient vertebrate speciation and local endemism in this region. Climate-driven rearrangements of drainage networks unrelated to tectonic activity contributed significantly to the assembly of species diversity and modern faunas in the East African biodiversity hotspot.

Eastern African environmental variation and its role in the evolution and cultural change of Homo over the last 1 million years.

Characterizing eastern African environmental variability on orbital timescales is crucial to evaluating the hominin evolutionary response to past climate changes. However, there is a dearth of high-resolution, well-dated records of ecosystem dynamics from eastern Africa that cover long time intervals. In the last 1 Myr, there were significant anatomical and cultural developments in Homo, including the origin of Homo sapiens. There were also major changes in global climatic boundary conditions that may have affected eastern African environments, yet potential linkages remain poorly understood. We developed carbon isotopic records from plant waxes (δ13Cwax) and bulk organic matter (δ13COM) from a well-dated sediment core spanning the last ∼1 Myr extracted from the Koora Basin, located south of the Olorgesailie Basin, in the southern Kenya rift. Our record characterizes the climatic and environmental context for evolutionary events and technological advances recorded in the adjacent Olorgesailie Basin, such as the transition from Acheulean to Middle Stone Age tools by 320 ka. A significant shift toward more C4-dominated ecosystems and arid conditions occurred near the end of the mid-Pleistocene Transition, which indicates a link between equatorial eastern African and high-latitude northern hemisphere climate. Environmental variability increases throughout the mid- to late-Pleistocene, superimposed by precession-paced packets of variability modulated by eccentricity. An interval of particularly high-amplitude climate and environmental variability occurred from ∼275 ka to ∼180 ka, synchronous with evidence for the first H. sapiens fossils in eastern Africa. These results support the 'variability selection hypothesis' that increased environmental variability selected for adaptable traits, behaviors, and technology in our hominin ancestors.

Increased ecological resource variability during a critical transition in hominin evolution.

Although climate change is considered to have been a large-scale driver of African human evolution, landscape-scale shifts in ecological resources that may have shaped novel hominin adaptations are rarely investigated. We use well-dated, high-resolution, drill-core datasets to understand ecological dynamics associated with a major adaptive transition in the archeological record ~24 km from the coring site. Outcrops preserve evidence of the replacement of Acheulean by Middle Stone Age (MSA) technological, cognitive, and social innovations between 500 and 300 thousand years (ka) ago, contemporaneous with large-scale taxonomic and adaptive turnover in mammal herbivores. Beginning ~400 ka ago, tectonic, hydrological, and ecological changes combined to disrupt a relatively stable resource base, prompting fluctuations of increasing magnitude in freshwater availability, grassland communities, and woody plant cover. Interaction of these factors offers a resource-oriented hypothesis for the evolutionary success of MSA adaptations, which likely contributed to the ecological flexibility typical of Homo sapiens foragers.

Widespread global peatland establishment and persistence over the last 130,000 y.

Glacial-interglacial variations in CO2 and methane in polar ice cores have been attributed, in part, to changes in global wetland extent, but the wetland distribution before the Last Glacial Maximum (LGM, 21 ka to 18 ka) remains virtually unknown. We present a study of global peatland extent and carbon (C) stocks through the last glacial cycle (130 ka to present) using a newly compiled database of 1,063 detailed stratigraphic records of peat deposits buried by mineral sediments, as well as a global peatland model. Quantitative agreement between modeling and observations shows extensive peat accumulation before the LGM in northern latitudes (>40°N), particularly during warmer periods including the last interglacial (130 ka to 116 ka, MIS 5e) and the interstadial (57 ka to 29 ka, MIS 3). During cooling periods of glacial advance and permafrost formation, the burial of northern peatlands by glaciers and mineral sediments decreased active peatland extent, thickness, and modeled C stocks by 70 to 90% from warmer times. Tropical peatland extent and C stocks show little temporal variation throughout the study period. While the increased burial of northern peats was correlated with cooling periods, the burial of tropical peat was predominately driven by changes in sea level and regional hydrology. Peat burial by mineral sediments represents a mechanism for long-term terrestrial C storage in the Earth system. These results show that northern peatlands accumulate significant C stocks during warmer times, indicating their potential for C sequestration during the warming Anthropocene.

Tropical peatland carbon storage linked to global latitudinal trends in peat recalcitrance.

Peatlands represent large terrestrial carbon banks. Given that most peat accumulates in boreal regions, where low temperatures and water saturation preserve organic matter, the existence of peat in (sub)tropical regions remains enigmatic. Here we examined peat and plant chemistry across a latitudinal transect from the Arctic to the tropics. Near-surface low-latitude peat has lower carbohydrate and greater aromatic content than near-surface high-latitude peat, creating a reduced oxidation state and resulting recalcitrance. This recalcitrance allows peat to persist in the (sub)tropics despite warm temperatures. Because we observed similar declines in carbohydrate content with depth in high-latitude peat, our data explain recent field-scale deep peat warming experiments in which catotelm (deeper) peat remained stable despite temperature increases up to 9 °C. We suggest that high-latitude deep peat reservoirs may be stabilized in the face of climate change by their ultimately lower carbohydrate and higher aromatic composition, similar to tropical peats.

A radiative forcing analysis of tropical peatlands before and after their conversion to agricultural plantations.

The tropical peat swamp forests of South-East Asia are being rapidly converted to agricultural plantations of oil palm and Acacia creating a significant global "hot-spot" for CO2 emissions. However, the effect of this major perturbation has yet to be quantified in terms of global warming potential (GWP) and the Earth's radiative budget. We used a GWP analysis and an impulse-response model of radiative forcing to quantify the climate forcing of this shift from a long-term carbon sink to a net source of greenhouse gases (CO2 and CH4 ). In the GWP analysis, five tropical peatlands were sinks in terms of their CO2 equivalent fluxes while they remained undisturbed. However, their drainage and conversion to oil palm and Acacia plantations produced a dramatic shift to very strong net CO2 -equivalent sources. The induced losses of peat carbon are ~20× greater than the natural CO2 sequestration rates. In contrast, a radiative forcing model indicates that the magnitude of this shift from a net cooling to warming effect is ultimately related to the size of an individual peatland's carbon pool. The continuous accumulation of carbon in pristine tropical peatlands produced a progressively negative radiative forcing (i.e., cooling) that ranged from -2.1 to -6.7 nW/m2 per hectare peatland by 2010 CE, referenced to zero at the time of peat initiation. Peatland conversion to plantations leads to an immediate shift from negative to positive trend in radiative forcing (i.e., warming). If drainage persists, peak warming ranges from +3.3 to +8.7 nW/m2 per hectare of drained peatland. More importantly, this net warming impact on the Earth's radiation budget will persist for centuries to millennia after all the peat has been oxidized to CO2 . This previously unreported and undesirable impact on the Earth's radiative balance provides a scientific rationale for conserving tropical peatlands in their pristine state.

How temporal patterns in rainfall determine the geomorphology and carbon fluxes of tropical peatlands.

Tropical peatlands now emit hundreds of megatons of carbon dioxide per year because of human disruption of the feedbacks that link peat accumulation and groundwater hydrology. However, no quantitative theory has existed for how patterns of carbon storage and release accompanying growth and subsidence of tropical peatlands are affected by climate and disturbance. Using comprehensive data from a pristine peatland in Brunei Darussalam, we show how rainfall and groundwater flow determine a shape parameter (the Laplacian of the peat surface elevation) that specifies, under a given rainfall regime, the ultimate, stable morphology, and hence carbon storage, of a tropical peatland within a network of rivers or canals. We find that peatlands reach their ultimate shape first at the edges of peat domes where they are bounded by rivers, so that the rate of carbon uptake accompanying their growth is proportional to the area of the still-growing dome interior. We use this model to study how tropical peatland carbon storage and fluxes are controlled by changes in climate, sea level, and drainage networks. We find that fluctuations in net precipitation on timescales from hours to years can reduce long-term peat accumulation. Our mathematical and numerical models can be used to predict long-term effects of changes in temporal rainfall patterns and drainage networks on tropical peatland geomorphology and carbon storage.

The evolutionary origin and population history of the grauer gorilla.

Gorillas living in western central Africa (Gorilla gorilla) are morphologically and genetically distinguishable from those living in eastern central Africa (Gorilla beringei). Genomic analyses show eastern gorillas experienced a significant reduction in population size during the Pleistocene subsequent to geographical isolation from their western counterparts. However, how these results relate more specifically to the recent biogeographical and evolutionary history of eastern gorillas remains poorly understood. Here we show that two rare morphological traits are present in the hands and feet of both eastern gorilla subspecies at strikingly high frequencies (>60% in G. b. graueri; ∼28% in G. b. beringei) in comparison with western gorillas (<1%). The intrageneric distribution of these rare traits suggests that they became common among eastern gorillas after diverging from their western relatives during the early to middle Pleistocene. The extremely high frequencies observed among grauer gorillas-which currently occupy a geographic range more than ten times the size of that of mountain gorillas-imply that grauers originated relatively recently from a small founding population of eastern gorillas. Current paleoenvironmental, geological, and biogeographical evidence supports the hypothesis that a small group of eastern gorillas likely dispersed westward from the Virungas into present-day grauer range in the highlands just north of Lake Kivu, either immediately before or directly after the Younger Dryas interval. We propose that as the lowland forests of central Africa expanded rapidly during the early Holocene, they became connected with the expanding highland forests along the Albertine Rift and enabled the descendants of this small group to widely disperse. The descendant populations significantly expanded their geographic range and population numbers relative to the gorillas of the Virunga Mountains and the Bwindi-Impenetrable Forest, ultimately resulting in the grauer gorilla subspecies recognized today. This founder-effect hypothesis offers some optimism for modern conservation efforts to save critically endangered eastern gorillas from extinction.