A biogeographical appraisal of the threatened South East Africa Montane Archipelago ecoregion.

Recent biological surveys of ancient inselbergs in southern Malawi and northern Mozambique have led to the discovery and description of many species new to science, and overlapping centres of endemism across multiple taxa. Combining these endemic taxa with data on geology and climate, we propose the 'South East Africa Montane Archipelago' (SEAMA) as a distinct ecoregion of global biological importance. The ecoregion encompasses 30 granitic inselbergs reaching > 1000 m above sea level, hosting the largest (Mt Mabu) and smallest (Mt Lico) mid-elevation rainforests in southern Africa, as well as biologically unique montane grasslands. Endemic taxa include 127 plants, 45 vertebrates (amphibians, reptiles, birds, mammals) and 45 invertebrate species (butterflies, freshwater crabs), and two endemic genera of plants and reptiles. Existing dated phylogenies of endemic animal lineages suggests this endemism arose from divergence events coinciding with repeated isolation of these mountains from the pan-African forests, together with the mountains' great age and relative climatic stability. Since 2000, the SEAMA has lost 18% of its primary humid forest cover (up to 43% in some sites)-one of the highest deforestation rates in Africa. Urgently rectifying this situation, while addressing the resource needs of local communities, is a global priority for biodiversity conservation.

Nature Forest Reserves in Tanzania and their importance for conservation.

Since 1997 Tanzania has undertaken a process to identify and declare a network of Nature Forest Reserves (NFRs) with high biodiversity values, from within its existing portfolio of national Forest Reserves, with 16 new NFRs declared since 2015. The current network of 22 gazetted NFRs covered 948,871 hectares in 2023. NFRs now cover a range of Tanzanian habitat types, including all main forest types-wet, seasonal, and dry-as well as wetlands and grasslands. NFRs contain at least 178 of Tanzania's 242 endemic vertebrate species, of which at least 50% are threatened with extinction, and 553 Tanzanian endemic plant taxa (species, subspecies, and varieties), of which at least 50% are threatened. NFRs also support 41 single-site endemic vertebrate species and 76 single-site endemic plant taxa. Time series analysis of management effectiveness tracking tool (METT) data shows that NFR management effectiveness is increasing, especially where donor funds have been available. Improved management and investment have resulted in measurable reductions of some critical threats in NFRs. Still, ongoing challenges remain to fully contain issues of illegal logging, charcoal production, firewood, pole-cutting, illegal hunting and snaring of birds and mammals, fire, wildlife trade, and the unpredictable impacts of climate change. Increased tourism, diversified revenue generation and investment schemes, involving communities in management, and stepping up control measures for remaining threats are all required to create a network of economically self-sustaining NFRs able to conserve critical biodiversity values.

Implications of zero-deforestation palm oil for tropical grassy and dry forest biodiversity.

Many companies have made zero-deforestation commitments (ZDCs) to reduce carbon emissions and biodiversity losses linked to tropical commodities. However, ZDCs conserve areas primarily based on tree cover and aboveground carbon, potentially leading to the unintended consequence that agricultural expansion could be encouraged in biomes outside tropical rainforest, which also support important biodiversity. We examine locations suitable for zero-deforestation expansion of commercial oil palm, which is increasingly expanding outside the tropical rainforest biome, by generating empirical models of global suitability for rainfed and irrigated oil palm. We find that tropical grassy and dry forest biomes contain >50% of the total area of land climatically suitable for rainfed oil palm expansion in compliance with ZDCs (following the High Carbon Stock Approach; in locations outside urban areas and cropland), and that irrigation could double the area suitable for expansion in these biomes. Within these biomes, ZDCs fail to protect areas of high vertebrate richness from oil palm expansion. To prevent unintended consequences of ZDCs and minimize the environmental impacts of oil palm expansion, policies and governance for sustainable development and conservation must expand focus from rainforests to all tropical biomes.

A practice-led assessment of landscape restoration potential in a biodiversity hotspot.

Effective restoration planning tools are needed to mitigate global carbon and biodiversity crises. Published spatial assessments of restoration potential are often at large scales or coarse resolutions inappropriate for local action. Using a Tanzanian case study, we introduce a systematic approach to inform landscape restoration planning, estimating spatial variation in cost-effectiveness, based on restoration method, logistics, biomass modelling and uncertainty mapping. We found potential for biomass recovery across 77.7% of a 53 000 km2 region, but with some natural spatial discontinuity in moist forest biomass, that was previously assigned to human causes. Most areas with biomass deficit (80.5%) were restorable through passive or assisted natural regeneration. However, cumulative biomass gains from planting outweighed initially high implementation costs meaning that, where applicable, this method yielded greater long-term returns on investment. Accounting for ecological, funding and other uncertainty, the top 25% consistently cost-effective sites were within protected areas and/or moderately degraded moist forest and savanna. Agro-ecological mosaics had high biomass deficit but little cost-effective restoration potential. Socio-economic research will be needed to inform action towards environmental and human development goals in these areas. Our results highlight value in long-term landscape restoration investments and separate treatment of savannas and forests. Furthermore, they contradict previously asserted low restoration potential in East Africa, emphasizing the importance of our regional approach for identifying restoration opportunities across the tropics. This article is part of the theme issue 'Understanding forest landscape restoration: reinforcing scientific foundations for the UN Decade on Ecosystem Restoration'.

High aboveground carbon stock of African tropical montane forests.

Tropical forests store 40-50 per cent of terrestrial vegetation carbon1. However, spatial variations in aboveground live tree biomass carbon (AGC) stocks remain poorly understood, in particular in tropical montane forests2. Owing to climatic and soil changes with increasing elevation3, AGC stocks are lower in tropical montane forests compared with lowland forests2. Here we assemble and analyse a dataset of structurally intact old-growth forests (AfriMont) spanning 44 montane sites in 12 African countries. We find that montane sites in the AfriMont plot network have a mean AGC stock of 149.4 megagrams of carbon per hectare (95% confidence interval 137.1-164.2), which is comparable to lowland forests in the African Tropical Rainforest Observation Network4 and about 70 per cent and 32 per cent higher than averages from plot networks in montane2,5,6 and lowland7 forests in the Neotropics, respectively. Notably, our results are two-thirds higher than the Intergovernmental Panel on Climate Change default values for these forests in Africa8. We find that the low stem density and high abundance of large trees of African lowland forests4 is mirrored in the montane forests sampled. This carbon store is endangered: we estimate that 0.8 million hectares of old-growth African montane forest have been lost since 2000. We provide country-specific montane forest AGC stock estimates modelled from our plot network to help to guide forest conservation and reforestation interventions. Our findings highlight the need for conserving these biodiverse9,10 and carbon-rich ecosystems.

Habitat availability explains variation in climate-driven range shifts across multiple taxonomic groups.

Range shifting is vital for species persistence, but there is little consensus on why individual species vary so greatly in the rates at which their ranges have shifted in response to recent climate warming. Here, using 40 years of distribution data for 291 species from 13 invertebrate taxa in Britain, we show that interactions between habitat availability and exposure to climate change at the range margins explain up to half of the variation in rates of range shift. Habitat generalists expanded faster than more specialised species, but this intrinsic trait explains less of the variation in range shifts than habitat availability, which additionally depends on extrinsic factors that may be rare or widespread at the range margin. Similarly, while climate change likely underlies polewards expansions, we find that more of the between-species variation is explained by differences in habitat availability than by changes in climatic suitability. A model that includes both habitat and climate, and their statistical interaction, explains the most variation in range shifts. We conclude that climate-change vulnerability assessments should focus as much on future habitat availability as on climate sensitivity and exposure, with the expectation that habitat restoration and protection will substantially improve species' abilities to respond to uncertain future climates.

Climate-induced phenology shifts linked to range expansions in species with multiple reproductive cycles per year.

Advances in phenology (the annual timing of species' life-cycles) in response to climate change are generally viewed as bioindicators of climate change, but have not been considered as predictors of range expansions. Here, we show that phenology advances combine with the number of reproductive cycles per year (voltinism) to shape abundance and distribution trends in 130 species of British Lepidoptera, in response to ~0.5 °C spring-temperature warming between 1995 and 2014. Early adult emergence in warm years resulted in increased within- and between-year population growth for species with multiple reproductive cycles per year (n = 39 multivoltine species). By contrast, early emergence had neutral or negative consequences for species with a single annual reproductive cycle (n = 91 univoltine species), depending on habitat specialisation. We conclude that phenology advances facilitate polewards range expansions in species exhibiting plasticity for both phenology and voltinism, but may inhibit expansion by less flexible species.

Correction to: Quantifying and understanding carbon storage and sequestration within the Eastern Arc Mountains of Tanzania, a tropical biodiversity hotspot.

Upon publication of the original article [1], the authors noticed that the figure labelling for Fig. 4 in the online version was processed wrong. The top left panel should be panel a, with the panels to its right being b and c. d and e should be the panels on the lower row, and f is correct. The graphs themselves are all correct. It is simply the letter labels that are wrong.

Climate change, climatic variation and extreme biological responses.

Extreme climatic events could be major drivers of biodiversity change, but it is unclear whether extreme biological changes are (i) individualistic (species- or group-specific), (ii) commonly associated with unusual climatic events and/or (iii) important determinants of long-term population trends. Using population time series for 238 widespread species (207 Lepidoptera and 31 birds) in England since 1968, we found that population 'crashes' (outliers in terms of species' year-to-year population changes) were 46% more frequent than population 'explosions'. (i) Every year, at least three species experienced extreme changes in population size, and in 41 of the 44 years considered, some species experienced population crashes while others simultaneously experienced population explosions. This suggests that, even within the same broad taxonomic groups, species are exhibiting individualistic dynamics, most probably driven by their responses to different, short-term events associated with climatic variability. (ii) Six out of 44 years showed a significant excess of species experiencing extreme population changes (5 years for Lepidoptera, 1 for birds). These 'consensus years' were associated with climatically extreme years, consistent with a link between extreme population responses and climatic variability, although not all climatically extreme years generated excess numbers of extreme population responses. (iii) Links between extreme population changes and long-term population trends were absent in Lepidoptera and modest (but significant) in birds. We conclude that extreme biological responses are individualistic, in the sense that the extreme population changes of most species are taking place in different years, and that long-term trends of widespread species have not, to date, been dominated by these extreme changes.This article is part of the themed issue 'Behavioural, ecological and evolutionary responses to extreme climatic events'.

Land cover change and carbon emissions over 100 years in an African biodiversity hotspot.

Agricultural expansion has resulted in both land use and land cover change (LULCC) across the tropics. However, the spatial and temporal patterns of such change and their resulting impacts are poorly understood, particularly for the presatellite era. Here, we quantify the LULCC history across the 33.9 million ha watershed of Tanzania's Eastern Arc Mountains, using geo-referenced and digitized historical land cover maps (dated 1908, 1923, 1949 and 2000). Our time series from this biodiversity hotspot shows that forest and savanna area both declined, by 74% (2.8 million ha) and 10% (2.9 million ha), respectively, between 1908 and 2000. This vegetation was replaced by a fivefold increase in cropland, from 1.2 million ha to 6.7 million ha. This LULCC implies a committed release of 0.9 Pg C (95% CI: 0.4-1.5) across the watershed for the same period, equivalent to 0.3 Mg C ha(-1)  yr(-1) . This is at least threefold higher than previous estimates from global models for the same study area. We then used the LULCC data from before and after protected area creation, as well as from areas where no protection was established, to analyse the effectiveness of legal protection on land cover change despite the underlying spatial variation in protected areas. We found that, between 1949 and 2000, forest expanded within legally protected areas, resulting in carbon uptake of 4.8 (3.8-5.7) Mg C ha(-1) , compared to a committed loss of 11.9 (7.2-16.6) Mg C ha(-1) within areas lacking such protection. Furthermore, for nine protected areas where LULCC data are available prior to and following establishment, we show that protection reduces deforestation rates by 150% relative to unprotected portions of the watershed. Our results highlight that considerable LULCC occurred prior to the satellite era, thus other data sources are required to better understand long-term land cover trends in the tropics.

Interactions between Canopy Structure and Herbaceous Biomass along Environmental Gradients in Moist Forest and Dry Miombo Woodland of Tanzania.

We have limited understanding of how tropical canopy foliage varies along environmental gradients, and how this may in turn affect forest processes and functions. Here, we analyse the relationships between canopy leaf area index (LAI) and above ground herbaceous biomass (AGBH) along environmental gradients in a moist forest and miombo woodland in Tanzania. We recorded canopy structure and herbaceous biomass in 100 permanent vegetation plots (20 m × 40 m), stratified by elevation. We quantified tree species richness, evenness, Shannon diversity and predominant height as measures of structural variability, and disturbance (tree stumps), soil nutrients and elevation as indicators of environmental variability. Moist forest and miombo woodland differed substantially with respect to nearly all variables tested. Both structural and environmental variables were found to affect LAI and AGBH, the latter being additionally dependent on LAI in moist forest but not in miombo, where other factors are limiting. Combining structural and environmental predictors yielded the most powerful models. In moist forest, they explained 76% and 25% of deviance in LAI and AGBH, respectively. In miombo woodland, they explained 82% and 45% of deviance in LAI and AGBH. In moist forest, LAI increased non-linearly with predominant height and linearly with tree richness, and decreased with soil nitrogen except under high disturbance. Miombo woodland LAI increased linearly with stem density, soil phosphorous and nitrogen, and decreased linearly with tree species evenness. AGBH in moist forest decreased with LAI at lower elevations whilst increasing slightly at higher elevations. AGBH in miombo woodland increased linearly with soil nitrogen and soil pH. Overall, moist forest plots had denser canopies and lower AGBH compared with miombo plots. Further field studies are encouraged, to disentangle the direct influence of LAI on AGBH from complex interrelationships between stand structure, environmental gradients and disturbance in African forests and woodlands.

Quantifying and understanding carbon storage and sequestration within the Eastern Arc Mountains of Tanzania, a tropical biodiversity hotspot.

BACKGROUND: The carbon stored in vegetation varies across tropical landscapes due to a complex mix of climatic and edaphic variables, as well as direct human interventions such as deforestation and forest degradation. Mapping and monitoring this variation is essential if policy developments such as REDD+ (Reducing Emissions from Deforestation and Forest Degradation) are to be known to have succeeded or failed. RESULTS: We produce a map of carbon storage across the watershed of the Tanzanian Eastern Arc Mountains (33.9 million ha) using 1,611 forest inventory plots, and correlations with associated climate, soil and disturbance data. As expected, tropical forest stores more carbon per hectare (182 Mg C ha(-1)) than woody savanna (51 Mg C ha(-1)). However, woody savanna is the largest aggregate carbon store, with 0.49 Pg C over 9.6 million ha. We estimate the whole landscape stores 1.3 Pg C, significantly higher than most previous estimates for the region. The 95% Confidence Interval for this method (0.9 to 3.2 Pg C) is larger than simpler look-up table methods (1.5 to 1.6 Pg C), suggesting simpler methods may underestimate uncertainty. Using a small number of inventory plots with two censuses (n = 43) to assess changes in carbon storage, and applying the same mapping procedures, we found that carbon storage in the tree-dominated ecosystems has decreased, though not significantly, at a mean rate of 1.47 Mg C ha(-1) yr(-1) (c. 2% of the stocks of carbon per year). CONCLUSIONS: The most influential variables on carbon storage in the region are anthropogenic, particularly historical logging, as noted by the largest coefficient of explanatory variable on the response variable. Of the non-anthropogenic factors, a negative correlation with air temperature and a positive correlation with water availability dominate, having smaller p-values than historical logging but also smaller influence. High carbon storage is typically found far from the commercial capital, in locations with a low monthly temperature range, without a strong dry season, and in areas that have not suffered from historical logging. The results imply that policy interventions could retain carbon stored in vegetation and likely successfully slow or reverse carbon emissions.

Towards regional, error-bounded landscape carbon storage estimates for data-deficient areas of the world.

Monitoring landscape carbon storage is critical for supporting and validating climate change mitigation policies. These may be aimed at reducing deforestation and degradation, or increasing terrestrial carbon storage at local, regional and global levels. However, due to data-deficiencies, default global carbon storage values for given land cover types such as 'lowland tropical forest' are often used, termed 'Tier 1 type' analyses by the Intergovernmental Panel on Climate Change (IPCC). Such estimates may be erroneous when used at regional scales. Furthermore uncertainty assessments are rarely provided leading to estimates of land cover change carbon fluxes of unknown precision which may undermine efforts to properly evaluate land cover policies aimed at altering land cover dynamics. Here, we present a repeatable method to estimate carbon storage values and associated 95% confidence intervals (CI) for all five IPCC carbon pools (aboveground live carbon, litter, coarse woody debris, belowground live carbon and soil carbon) for data-deficient regions, using a combination of existing inventory data and systematic literature searches, weighted to ensure the final values are regionally specific. The method meets the IPCC 'Tier 2' reporting standard. We use this method to estimate carbon storage over an area of33.9 million hectares of eastern Tanzania, reporting values for 30 land cover types. We estimate that this area stored 6.33 (5.92-6.74) Pg C in the year 2000. Carbon storage estimates for the same study area extracted from five published Africa-wide or global studies show a mean carbon storage value of ∼50% of that reported using our regional values, with four of the five studies reporting lower carbon storage values. This suggests that carbon storage may have been underestimated for this region of Africa. Our study demonstrates the importance of obtaining regionally appropriate carbon storage estimates, and shows how such values can be produced for a relatively low investment.

Protected areas: mixed success in conserving East Africa's evergreen forests.

In East Africa, human population growth and demands for natural resources cause forest loss contributing to increased carbon emissions and reduced biodiversity. Protected Areas (PAs) are intended to conserve habitats and species. Variability in PA effectiveness and 'leakage' (here defined as displacement of deforestation) may lead to different trends in forest loss within, and adjacent to, existing PAs. Here, we quantify spatial variation in trends of evergreen forest coverage in East Africa between 2001 and 2009, and test for correlations with forest accessibility and environmental drivers. We investigate PA effectiveness at local, landscape and national scales, comparing rates of deforestation within park boundaries with those detected in park buffer zones and in unprotected land more generally. Background forest loss (BFL) was estimated at -9.3% (17,167 km(2)), but varied between countries (range: -0.9% to -85.7%; note: no BFL in South Sudan). We document high variability in PA effectiveness within and between PA categories. The most successful PAs were National Parks, although only 26 out of 48 parks increased or maintained their forest area (i.e. Effective parks). Forest Reserves (Ineffective parks, i.e. parks that lose forest from within boundaries: 204 out of 337), Nature Reserves (six out of 12) and Game Parks (24 out of 26) were more likely to lose forest cover. Forest loss in buffer zones around PAs exceeded background forest loss, in some areas indicating leakage driven by Effective National Parks. Human pressure, forest accessibility, protection status, distance to fires and long-term annual rainfall were highly significant drivers of forest loss in East Africa. Some of these factors can be addressed by adjusting park management. However, addressing close links between livelihoods, natural capital and poverty remains a fundamental challenge in East Africa's forest conservation efforts.

The species-area relationship and confounding variables in a threatened monkey community.

This study investigates the species-area relationship (SAR) for forest monkeys in a biodiversity hotspot. The Udzungwa Mountains of Tanzania are well-suited to investigate the SAR, with seven monkey species in a range of fragment sizes (0.06-526 km(2)). We test the relationship between species richness and forest fragment size, relative to human and environmental factors. We distinguish resident and transitory species because the latter have an "effective patch size" beyond the area of forest. Forest area was the strongest (log-linear) predictor of species richness. However, forest area, elevation range and annual moisture index were intercorrelated. Previous knowledge of the relationship between elevation and tree communities suggests that the SAR is largely a result of habitat heterogeneity. Isolation by farmland (matrix habitat) also had a significant negative effect on species richness, probably exacerbated by hunting in small forests. The effect of area and isolation was less for transitory species. The human influence on species' presence/absence was negatively related to the extent of occurrence. Weaker relationships with temperature and precipitation suggest underlying climatic influences, and give some support for the influence of productivity. A reduced area relationship for smaller forests suggests that fragment sizes below 12-40 km(2) may not be reliable for determining SAR in forest monkeys. Further practical implications are for management to encourage connectivity, and for future SAR research to consider residency, matrix classification and moisture besides precipitation.