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Nonparametric methods are widely used for assessing the performance of firms and other organizations in the private and public sectors. Typically, FDH or DEA estimators that estimate the attainable sets and its efficient boundary by enveloping the cloud of observed units in the appropriate input-output space are used. The statistical properties of these estimators have been established and inference is available using appropriate nonparametric techniques. In particular, hypotheses on model structure and the production process can be tested using using recent theoretical results including Central Limit Theorems on limiting distribution of means of efficiency scores. This chapter shows how these results can be used for testing the equality of means of efficiency, convexity of production sets and separability with respect to environmental factors are considered, and in addition for analyzing the dynamics of the production process over time. An empirical illustration is provided by using the various results and tests to examine the performance of municipal governments in the U.S. in providing local public goods.

Woodland caribou ( Rangifer tarandus caribou) are threatened in Canada because of the drastic decline in population size caused primarily by human-induced landscape changes that decrease habitat and increase predation risk. Conservation efforts have largely focused on reducing predators and protecting critical habitat, whereas research on dietary niches and the role of potential food constraints in lichen-poor environments is limited. To improve our understanding of dietary niche variability, we used a next-generation sequencing approach with metabarcoding of DNA extracted from faecal pellets of woodland caribou located on Lake Superior in lichen-rich (mainland) and lichen-poor (island) environments. Amplicon sequencing of fungal ITS2 region revealed lichen-associated fungi as predominant in samples from both populations, but amplification at the chloroplast trnL region, which was only successful on island samples, revealed primary consumption of yew ( Taxus spp.) based on relative read abundance (83.68%) with dogwood ( Cornus spp.; 9.67%) and maple ( Acer spp.; 4.10%) also prevalent. These results suggest that conservation efforts for caribou need to consider the availability of food resources beyond lichen to ensure successful outcomes. More broadly, we provide a reliable methodology for assessing ungulate diet from archived faecal pellets that could reveal important dietary shifts over time in response to climate change.

This paper estimates the shadow price of equity for U.S. commercial banks over 2001–2018 using nonparametric local-linear estimators of the underlying cost frontier and tests the existence of “Too-Big-to-Fail” (TBTF) banks. Evidence for the existence of TBTF banks is found. We find that a negative correlation exists between the shadow price of equity and the size of banks in each year, suggesting that big banks pay less for equity than small banks. In addition, in each year there are more banks with a negative shadow price of equity in the fourth quartile based on total assets than in the other three quartiles. The data also reveal that for each year, the estimated mean shadow price of equity for the 50 largest banks is smaller than the mean price of deposits, even though equity is commonly viewed as a riskier asset than deposits. Finally, we find that the top 10 largest banks are willing to pay much more at the start of the global financial crisis and after the Dodd-Frank Wall Street Reform and Consumer Protection Act of 2010 than the other periods. These results imply that these regulations are effective in reducing the implicit subsidy, at least for the top 10 largest banks. However, it is also evident that the recapitalization has imposed significant equity funding costs for the top 10 largest banks.

Intra-specific diversification in the northern hemisphere is a recent phenomenon, particularly for vertebrates, due to climatic oscillations during the Quaternary. Caribou (Rangifer tarandus) are a recently diversified, keystone species in North America with a continental-wide distribution and a large amount of phenotypic variation. We assess patterns of intra-specific genetic variation and adaptive divergence, which are of critical interest to the fields of evolutionary biology and conservation. We characterized the relationship among caribou ecotypes and rapidly evolving genes among intra-specific lineages using phylogenomic analyses and a newly assembled chromosome-scale reference genome and 66 high-coverage genomes. Nine phylogenomic lineages showed different levels of introgression, with some lineages highly introgressed. Despite this, dN/dS ratios of all genes in the caribou genome (>30,000) identified 700-800 genes with significant signatures of positive selection in each lineage. These genes function in diverse pathways ranging from immune responses to circulatory systems, with the most enriched molecular pathways found in lineages with high levels of introgression. Glacial cycles and repeated introgression events between caribou lineages likely inflated both neutral and adaptive genetic variation: findings that are particularly timely given the importance of intra-specific diversity and adaptive variation in wild species for resilience to global change.

Density is an important demographic parameter that is commonly overlooked in studies of wild populations. Here, we examined the effects of variable spatially explicit density on a range of demographic parameters in a wild population of a cryptic ungulate, boreal woodland caribou (Rangifer tarandus caribou). Using non-invasive genetic sampling, we applied spatial capture–recapture methods with landscape covariates to estimate the density of boreal woodland caribou across a 108,806 km2 study area. We then created a familial network from the reconstructed parent–offspring relationships to determine whether spatial density influenced sex-specific individual reproductive success, female pregnancy status, and dispersal distance. We found that animal density varied greatly in response to land cover types and disturbance; animal density was most influenced by landscape composition and distance to roads varying from 0 in areas with >20% deciduous cover to 270 caribou per 1,000 km2 in areas presenting contiguous older coniferous cover. We found that both male and female reproductive success varied with density, with males showing a higher probability of having offspring in higher-density areas, and the opposite for females. No differences were found in female pregnancy rates occurring in high- and low-density areas. Dispersal distances varied with density, with offspring moving shorter distances when parents were found in higher-density areas. Familial networks showed lower-closeness centrality and lower-degree centrality for females in higher-density areas, indicating that females found in higher-density areas tend to be less broadly associated with animals across the range. Although high-density areas do reflect good-quality caribou habitat, the observed decreased closeness and degree centrality measures, dispersal rates, and lower female recruitment rates suggest that remnant habitat patches across the landscape may create population sinks.

Conservation genomics is an important tool to manage threatened species under current biodiversity loss. Recent advances in sequencing technology mean that we can now use whole genomes to investigate demographic history, local adaptation, inbreeding, and more in unprecedented detail. However, for many rare and elusive species only non-invasive samples such as faeces can be obtained, making it difficult to take advantage of whole genome data. We present a method to extract DNA from the mucosal layer of faecal samples to re-sequence high coverage whole genomes using standard laboratory techniques. We use wild collected faecal pellets collected from caribou (Rangifer tarandus), a species undergoing declines in many parts of its range in Canada and subject to comprehensive conservation and population monitoring measures. We compare four faecal genomes to two tissue genomes sequenced in the same run. Quality metrics were similar between faecal and tissue samples with the main difference being the alignment success of raw reads to the reference genome due to differences in low quality and endogenous DNA content, affecting overall coverage. One of our faecal genomes was only re-sequenced at low coverage (1.6 ×), however the other three obtained between 7 and 15 ×, compared to 19 and 25 × for the tissue samples. We successfully re-sequenced high-quality whole genomes from faecal DNA and are one of the first to obtain genome-wide data from wildlife faecal DNA in a non-primate species. Our work represents an important advancement for non-invasive conservation genomics.

Pleistocene glacial cycles influenced the diversification of high-latitude wildlife species through recurrent periods of range contraction, isolation, divergence, and expansion from refugia and subsequent admixture of refugial populations. We investigate population size changes and the introgressive history of caribou (Rangifer tarandus) in western Canada using 33 whole genome sequences coupled with larger-scale mitochondrial data. We found that a major population expansion of caribou occurred starting around 110,000 years ago (kya), the start of the last glacial period. Additionally, we found effective population sizes of some caribou reaching ~700,000 to 1,000,000 individuals, one of the highest recorded historical effective population sizes for any mammal species thus far. Mitochondrial analyses dated introgression events prior to the LGM dating to 20-30 kya and even more ancient at 60 kya, coinciding with colder periods with extensive ice coverage, further demonstrating the importance of glacial cycles and events prior to the LGM in shaping demographic history. Reconstructing the origins and differential introgressive history has implications for predictions on species responses under climate change. Our results have implications for other whole genome analyses using pairwise sequentially Markovian coalescent (PSMC) analyses, as well as highlighting the need to investigate pre-LGM demographic patterns to fully reconstruct the origin of species diversity, especially for high-latitude species.

The evolutionary origins and hybridization patterns of Canis species in North America have been hotly debated for the past 30 years. Disentangling ancestry and timing of hybridization in Great Lakes wolves, eastern Canadian wolves, red wolves, and eastern coyotes are most often partitioned into a 2‐species model that assigns all ancestry to gray wolves and/or coyotes, and a 3‐species model that includes a third, North American evolved eastern wolf genome. The proposed models address recent or sometimes late Holocene hybridization events but have largely ignored potential Pleistocene era progenitors and opportunities for hybridization that may have impacted the current mixed genomes in eastern Canada and the United States. Here, we re‐analyze contemporary and ancient mitochondrial DNA genomes with Bayesian phylogenetic analyses to more accurately estimate divergence dates among lineages. We combine that with a review of the literature on Late Pleistocene Canis distributions to: (a) identify potential Pleistocene progenitors to southern North American gray wolves and eastern wolves; and (b) illuminate opportunities for ancient hybridization events. Specifically, we propose that Beringian gray wolves (C. lupus) and extinct large wolf‐like coyotes (C. latrans orcutti) are likely progenitors to Mexican and Plains gray wolves and eastern wolves, respectively, and may represent a potentially unrecognized source of introgressed genomic variation within contemporary Canis genomes. These events speak to the potential origins of contemporary genomes and provide a new perspective on Canis ancestry, but do not negate current conservation priorities of dwindling wolf populations with unique genomic signatures and key ecologically critical roles.

The Sahara is the largest hot desert on Earth. Yet the timing of its inception and its response to climatic forcing is debated, leading to uncertainty over the causes and consequences of regional aridity. Here we present detailed records of terrestrial inputs from Africa to North Atlantic deep-sea sediments, documenting a long and sustained history of astronomically paced oscillations between a humid and arid Sahara from over 11 million years ago. We show that intervals of strong dust emissions from the heart of the continent predate both the intensification of Northern Hemisphere glaciation and the oldest land-based evidence for a Saharan desert by millions of years. We find no simple long-term gradational transition towards an increasingly arid climate state in northern Africa, suggesting that aridity was not the primary driver of gradual Neogene expansion of African savannah C4 grasslands. Instead, insolation-driven wet–dry shifts in Saharan climate were common over the past 11 Myr, and we identify three distinct stages in the sensitivity of this relationship. Our data provide context for evolutionary outcomes on Africa; for example, we find that astronomically paced arid intervals predate the oldest fossil evidence of hominid bipedalism by at least 4 Myr.

Past hydroclimate change in Africa and the Arabian Peninsula plays a vital role in early human evolution and dispersal. Although arid today, North Africa and the Arabian Peninsula experienced periodic humid intervals, during which strengthened rainfall driven by astronomical forcing (chiefly eccentricity-modulated precessional cycles) transformed the Saharo-Arabian desert into a vegetated landscape cross-cut by rivers and lakes. The relationship between hydroclimate variability and hominid evolution is strongly debated and continuous palaeoclimate reconstructions are required to provide a context for understanding evolutionary outcomes on land. However, there are major discrepancies between different types of proxy data and between these reconstructions and those generated by numerical models as to the spatial and temporal occurrence of humid events and to the seasonality of the rainfall responsible. Two major contributing problems concern (i) the incontinuous nature of palaeo records from terrestrial archives (palaeolake sediments and stalagmites) and (ii) the challenges of attribution associated with records from marine sediment cores. Here, we report geochemical and stable isotope data from the Arabian Sea to reconstruct past environmental changes in a multi-proxy approach, which will help us to understand the relationship between palaeoclimate change and hominid evolution, reveal how the West Asian monsoon responded to a period of increased global warmth and furthermore, provide a reliable data set for model simulations in the future.

Highlights • We present LGC record of the Pb isotope composition Labrador Sea seawater. • These data can be used to track Laurentide Ice Sheet extent over Hudson Bay. • LIS retreat during the PGM was relatively fast compared to the LGM • The LIS first advanced significantly over Hudson Bay during MIS 4. • Our record does not support significant LIS retreat during MIS 3. Understanding the history of continental ice-sheet growth on North America, and in particular that of the Laurentide Ice Sheet (LIS), is important for palaeoclimate and sea-level reconstructions. Information on ice-sheet extent pre-dating the Last Glacial Maximum (LGM) is heavily reliant, though, on the outputs of numerical models underpinned by scant geological data. Important aspects of LIS history that remain unresolved include the timing of its collapse during Termination 2, the first time that it expanded significantly during the Last Glacial Cycle, and whether its volume was significantly reduced during marine isotope stage (MIS) 3. To address these issues and more, we present authigenic iron-manganese (Fe–Mn) oxyhydroxide-derived high-resolution records of Pb isotope data and associated rare earth element profiles for samples spanning the past ∼130 kyr from northwest North Atlantic Labrador Sea, IODP Site U1302/3. We use these new data to track chemical weathering intensity and solute flux to the Labrador Sea associated with LIS extent on the adjacent highly radiogenic (high Pb isotope composition) North American Superior Province (SP) craton since the Penultimate Glacial Maximum (PGM). Our new records show that relatively high (radiogenic) values characterise warm marine isotope stages (MIS) 5, 3 and 1 and the lowest (most unradiogenic) values occurred during cold stages MIS 6, 4 and 2. The radiogenic Pb isotope excursion associated with Termination 2 is short-lived relative to the one documented for Termination 1, suggesting that LIS retreat during the PGM was relatively fast compared to the LGM and that its collapse during the last interglacial occurred ∼125 ka. Highly radiogenic inputs to the Labrador Sea during MIS 5d-a, ∼116–71 ka, most likely reflect a spin-up in Labrador Current vigour, incipient glaciation and renewed glacial erosion of high grounds of the eastern SP craton by localised wet-based ice-caps. A large decrease in Pb isotope values towards unradiogenic LGM-like compositions between ∼75–65 ka across the MIS 5/4 transition likely reflects a slow-down in Labrador Current vigour, an increase in subaerial deposition of aeolian dust and a significant advance of the LIS across Hudson Bay caused a strong reduction or even abandonment of Pb sourcing from the SP. The relatively radiogenic Pb isotope composition of bottom-waters bathing our study site during MIS 3, 57–29 ka, is unlikely to support a recently proposed major reduction in LIS extent for this time. Instead, we argue these values are better explained by southern Greenland Ice Sheet retreat, increased chemical weathering of the Ketelidian Mobile Belt and subsequent Pb runoff from Greenland.

In social species, reproductive success and rates of dispersal vary among individuals resulting in spatially structured populations. Network analyses of familial relationships may provide insights on how these parameters influence population‐level demographic patterns. These methods, however, have rarely been applied to genetically derived pedigree data from wild populations.Here, we use parent–offspring relationships to construct familial networks from polygamous boreal woodland caribou (Rangifer tarandus caribou) in Saskatchewan, Canada, to inform recovery efforts. We collected samples from 933 individuals at 15 variable microsatellite loci along with caribou‐specific primers for sex identification. Using network measures, we assess the contribution of individual caribou to the population with several centrality measures and then determine which measures are best suited to inform on the population demographic structure. We investigate the centrality of individuals from eighteen different local areas, along with the entire population.We found substantial differences in centrality of individuals in different local areas, that in turn contributed differently to the full network, highlighting the importance of analyzing networks at different scales. The full network revealed that boreal caribou in Saskatchewan form a complex, interconnected familial network, as the removal of edges with high betweenness did not result in distinct subgroups. Alpha, betweenness, and eccentricity centrality were the most informative measures to characterize the population demographic structure and for spatially identifying areas of highest fitness levels and family cohesion across the range. We found varied levels of dispersal, fitness, and cohesion in family groups. Synthesis and applications: Our results demonstrate the value of different network measures in assessing genetically derived familial networks. The spatial application of the familial networks identified individuals presenting different fitness levels, short‐ and long‐distance dispersing ability across the range in support of population monitoring and recovery efforts., Network analyses of familial relationships may provide insights on how fitness influences population‐level demographic patterns in socially structured populations. Here, we use networks to reconstruct parent‐offspring relationships and create a familial network from polygamous boreal woodland caribou (Rangifer tarandus caribou), assessing the contribution of individual caribou to the population with several centrality metrics and determine which metrics are best suited to inform on population demographic structure. The familial network revealed that boreal caribou in Saskatchewan form a complex, interconnected social network, and identified areas of higher fitness levels and social cohesion across the range in support of population monitoring and recovery efforts.

Kalman filter-based retrodiction plays an indispensable role in modern multitarget tracking and retrodiction (MTTR) algorithms. To this end, the Rauch–Tung–Striebel (RTS) smoother is a widely used Kalman filter-based target state smoother. With a large number of targets, MTTR algorithms, particularly with large window sizes, become very computationally intensive. If not addressed, these algorithms will not meet the requirements for tracking a large number of targets in real time. A natural approach is to parallelize these algorithms to render them useful, particularly in the context of emerging multicore platforms. However, this is nontrivial, as the governing mathematical framework of the RTS smoother, namely the dependencies between complex computations, prevents any form of parallelization. Although the MTTR component can naively be parallelized ignoring the smoothing component, the overall benefit, as we demonstrate in this article, is a fraction of the best possible benefits. In this article, by carefully reformulating the underlying mathematical framework that is necessary for retrodiction, we propose a novel, easily parallelizable RTS smoother. The proposed parallelized RTS smoother we outline in this article has best data reuse and enables the overall MTTR problem to be parallelized more efficiently. We demonstrate this on a state-of-the-art multicore processor platform using the shared-memory parallelism. Our results show that the parallel MTTR solution, which includes gating, assignment, tracking, and retrodiction, can offer nearly 150 times speed up against a fully sequential version. With excellent computational performance, our proposed RTS smoother enables very large window sizes with little or no impact on the overall performance.

Clock genes exhibit substantial control over gene expression and ultimately life-histories using external cues such as photoperiod, and are thus likely to be critical for adaptation to shifting seasonal conditions and novel environments as species redistribute their ranges under climate change. Coding trinucleotide repeats (cTNRs) are found within several clock genes, and may be interesting targets of selection due to their containment within exonic regions and elevated mutation rates. Here, we conduct inter-specific characterization of the NR1D1 cTNR between Canada lynx and bobcat, and intra-specific spatial and environmental association analyses of neutral microsatellites and our functional cTNR marker, to investigate the role of selection on this locus in Canada lynx. We report signatures of divergent selection between lynx and bobcat, with the potential for hybrid-mediated gene flow in the area of range overlap. We also provide evidence that this locus is under selection across Canada lynx in eastern Canada, with both spatial and environmental variables significantly contributing to the explained variation, after controlling for neutral population structure. These results suggest that cTNRs may play an important role in the generation of functional diversity within some mammal species, and allow for contemporary rates of adaptation in wild populations in response to environmental change. We encourage continued investment into the study of cTNR markers to better understand their broader relevance to the evolution and adaptation of mammals.

Accurately estimating abundance is a critical component of monitoring and recovery of rare and elusive species. Spatial capture–recapture (SCR) models are an increasingly popular method for robust estimation of ecological parameters. We provide an analytical framework to assess results from empirical studies to inform SCR sampling design, using both simulated and empirical data from noninvasive genetic sampling of seven boreal caribou populations (Rangifer tarandus caribou), which varied in range size and estimated population density. We use simulated population data with varying levels of clustered distributions to quantify the impact of nonindependence of detections on density estimates, and empirical datasets to explore the influence of varied sampling intensity on the relative bias and precision of density estimates. Simulations revealed that clustered distributions of detections did not significantly impact relative bias or precision of density estimates. The genotyping success rate of our empirical dataset (n = 7,210 samples) was 95.1%, and 1,755 unique individuals were identified. Analysis of the empirical data indicated that reduced sampling intensity had a greater impact on density estimates in smaller ranges. The number of captures and spatial recaptures was strongly correlated with precision, but not absolute relative bias. The best sampling designs did not differ with estimated population density but differed between large and small ranges. We provide an efficient framework implemented in R to estimate the detection parameters required when designing SCR studies. The framework can be used when designing a monitoring program to minimize effort and cost while maximizing effectiveness, which is critical for informing wildlife management and conservation., We provide an analytical framework to assess results from empirical studies to inform spatial capture–recapture sampling design, using both simulated and empirical data from noninvasive genetic sampling of seven boreal caribou populations (Rangifer tarandus caribou). Simulations revealed that clustered distributions of detections did not significantly impact relative bias or precision of density estimates, and analysis of the empirical data indicated that reduced sampling intensity had a greater impact on density estimates in smaller ranges. The framework can be used when designing a monitoring program to minimize effort and cost while maximizing effectiveness, which is critical for informing wildlife management and conservation.

Glioblastoma multiforme (GBM) remains the top challenge to radiotherapy with only 25% one-year survival after diagnosis. Here, we reveal that co-enhancement of mitochondrial fatty acid oxidation (FAO) enzymes (CPT1A, CPT2 and ACAD9) and immune checkpoint CD47 is dominant in recurrent GBM patients with poor prognosis. A glycolysis-to-FAO metabolic rewiring is associated with CD47 anti-phagocytosis in radioresistant GBM cells and regrown GBM after radiation in syngeneic mice. Inhibition of FAO by CPT1 inhibitor etomoxir or CRISPR-generated CPT1A−/−, CPT2−/−, ACAD9−/− cells demonstrate that FAO-derived acetyl-CoA upregulates CD47 transcription via NF-κB/RelA acetylation. Blocking FAO impairs tumor growth and reduces CD47 anti-phagocytosis. Etomoxir combined with anti-CD47 antibody synergizes radiation control of regrown tumors with boosted macrophage phagocytosis. These results demonstrate that enhanced fat acid metabolism promotes aggressive growth of GBM with CD47-mediated immune evasion. The FAO-CD47 axis may be targeted to improve GBM control by eliminating the radioresistant phagocytosis-proofing tumor cells in GBM radioimmunotherapy.

(131)I-metaiodobenzylguanidine ((131)I-mIBG) is a targeted radiation therapy developed for the treatment of advanced neuroblastoma. We have previously shown that this patient cohort can be used to predict absorbed dose associated with early (131)I exposure, 72 h after treatment. We now expand these studies to identify gene expression differences associated with (131)I-mIBG exposure 15 days after treatment. Total RNA from peripheral blood lymphocytes was isolated from 288 whole blood samples representing 59 relapsed or refractory neuroblastoma patients before and after (131)I-mIBG treatment. We found that several transcripts predictive of early exposure returned to baseline levels by day 15, however, selected transcripts did not return to baseline. At 72 h, all 17 selected pathway-specific transcripts were differentially expressed. Transcripts CDKN1A (P < 0.000001), FDXR (P < 0.000001), DDB2 (P < 0.000001), and BBC3 (P < 0.000001) showed the highest up-regulation at 72 h after (131)I-mIBG exposure, with mean log(2) fold changes of 2.55, 2.93, 1.86 and 1.85, respectively. At day 15 after (131)I-mIBG, 11 of the 17 selected transcripts were differentially expressed, with XPC, STAT5B, PRKDC, MDM2, POLH, IGF1R, and SGK1 displaying significant up-regulation at 72 h and significant down-regulation at day 15. Interestingly, transcripts FDXR (P = 0.01), DDB2 (P = 0.03), BCL2 (P = 0.003), and SESN1 (P < 0.0003) maintained differential expression 15 days after (131)I-mIBG treatment. These results suggest that transcript levels for DNA repair, apoptosis, and ionizing radiation-induced cellular stress are still changing by 15 days after (131)I-mIBG treatment. Our studies showcase the use of biodosimetry gene expression panels as predictive biomarkers following early (72 h) and late (15 days) internal (131)I exposure. Our findings also demonstrate the utility of our transcript panel to differentiate exposed from non-exposed individuals up to 15 days after exposure from internal (131)I.

Many instructors globally reported a lack of engagement in synchronous online sessions during the Covid-19 lockdowns. This article outlines the use of collaborative small group tasks mediated via breakout rooms in Teams during the 2020/21 academic year. The rationale for the activities, which are available for download, is described along with details of the evaluation of their impact. Key findings were that a majority of students reported enjoying the tasks and felt that they improved their learning during online sessions.

Stochastic frontier models along the lines of Aigner et al. are widely used to benchmark firms’ performances in terms of efficiency. The models are typically fully parametric, with functional form specifications for the frontier as well as both the noise and the inefficiency processes. Studies such as Kumbhakar et al. have attempted to relax some of the restrictions in parametric models, but so far all such approaches are limited to a univariate response variable. Some (e.g., Simar and Zelenyuk; Kuosmanen and Johnson) have proposed nonparametric estimation of directional distance functions to handle multiple inputs and outputs, raising issues of endogeneity that are either ignored or addressed by imposing restrictive and implausible assumptions. This article extends nonparametric methods developed by Simar et al. and Hafner et al. to allow multiple inputs and outputs in an almost fully nonparametric framework while avoiding endogeneity problems. We discuss properties of the resulting estimators, and examine their finite-sample performance through Monte Carlo experiments. Practical implementation of the method is illustrated using data on U.S. commercial banks.

North Africa features some of the most frequently burnt biomes on Earth, including the semi-arid grasslands of the Sahel and wetter savannas immediately to the south. Natural fires are fuelled by rapid biomass production during the wet season, its desiccation during the dry season and ignition by frequent dry lightning strikes. Today, fire activity decreases markedly both to the north of the Sahel, where rainfall is extremely low, almost eliminating biomass over the Sahara, and to the south where forest biomes are too wet to burn. Over the last glacial cycle, rainfall and vegetation cover over northern Africa varied dramatically in response to gradual astronomically-forced insolation change, changes in atmospheric carbon dioxide levels, and abrupt cooling events over the North Atlantic Ocean associated with the reorganisation of Meridional Overturning Circulation (MOC). Here we report the results of a study into the impact of these climate changes on fire activity in northern African over the last 50,000 years (50 kyr). Our reconstructions come from marine sediments with strong age control that provide an uninterrupted record of charcoal particles exported from the African continent. We studied three sites on a latitudinal transect along the northwest African margin between 21 and 9°N. Our sites exhibit a distinct latitudinal relationship between past changes in rainfall and fire activity. At the southernmost site (GeoB9528-3, 9°N), fire activity decreased during intervals of increasing humidity, while our northernmost site (ODP Site 658, 21°N) clearly demonstrates the opposite relationship. The site in the middle of our transect, offshore of the present day southern Sahel today (GeoB9508-5, 15°N), exhibits a “Goldilocks” relationship between fire activity and hydroclimate, wherein charcoal fluxes peak under intermediate rainfall climate conditions and are supressed by transition to more arid or more humid conditions. Our results are remarkably consistent with the predictions of the intermediate fire-productivity hypothesis developed in conceptual macroecological models and supported by empirical evidence of modern day fire activity. Feedback processes operating between fire, climate and vegetation are undoubtedly complex but temperature is suggested to be the main driver of temporal change in fire activity globally, with the precipitation-evaporation balance perhaps a secondary influence in the Holocene tropics. However, there is only sparse coverage of Africa in the composite records upon which those interpretations are based. We conclude that hydroclimate (not temperature) exerted the dominant control on burning in the tropics of northern Africa well before the Holocene (from at least 50 ka). publishedVersion