Environmental context shapes the relationship between grass consumption and body size in African herbivore communities.

Though herbivore grass dependence has been shown to increase with body size across herbivore species, it is unclear whether this relationship holds at the community level. Here we evaluate whether grass consumption scales positively with body size within African large mammalian herbivore communities and how this relationship varies with environmental context. We used stable carbon isotope and community occurrence data to investigate how grass dependence scales with body size within 23 savanna herbivore communities throughout eastern and central Africa. We found that dietary grass fraction increased with body size for the majority of herbivore communities considered, especially when complete community data were available. However, the slope of this relationship varied, and rainfall seasonality and elephant presence were key drivers of the variation-grass dependence increased less strongly with body size where rainfall was more seasonal and where elephants were present. We found also that the dependence of the herbivore community as a whole on grass peaked at intermediate woody cover. Intraspecific diet variation contributed to these community-level patterns: common hippopotamus (Hippopotamus amphibius) and giraffe (Giraffa camelopardalis) ate less grass where rainfall was more seasonal, whereas Cape buffalo (Syncerus caffer) and savanna elephant (Loxodonta africana) grass consumption were parabolically related to woody cover. Our results indicate that general rules appear to govern herbivore community assembly, though some aspects of herbivore foraging behavior depend upon local environmental context.

Collapse of terrestrial mammal food webs since the Late Pleistocene.

Food webs influence ecosystem diversity and functioning. Contemporary defaunation has reduced food web complexity, but simplification caused by past defaunation is difficult to reconstruct given the sparse paleorecord of predator-prey interactions. We identified changes to terrestrial mammal food webs globally over the past ~130,000 years using extinct and extant mammal traits, geographic ranges, observed predator-prey interactions, and deep learning models. Food webs underwent steep regional declines in complexity through loss of food web links after the arrival and expansion of human populations. We estimate that defaunation has caused a 53% decline in food web links globally. Although extinctions explain much of this effect, range losses for extant species degraded food webs to a similar extent, highlighting the potential for food web restoration via extant species recovery.

Early Pleistocene large mammals from Maka'amitalu, Hadar, lower Awash Valley, Ethiopia.

The Early Pleistocene was a critical time period in the evolution of eastern African mammal faunas, but fossil assemblages sampling this interval are poorly known from Ethiopia's Afar Depression. Field work by the Hadar Research Project in the Busidima Formation exposures (~2.7-0.8 Ma) of Hadar in the lower Awash Valley, resulted in the recovery of an early Homo maxilla (A.L. 666-1) with associated stone tools and fauna from the Maka'amitalu basin in the 1990s. These assemblages are dated to ~2.35 Ma by the Bouroukie Tuff 3 (BKT-3). Continued work by the Hadar Research Project over the last two decades has greatly expanded the faunal collection. Here, we provide a comprehensive account of the Maka'amitalu large mammals (Artiodactyla, Carnivora, Perissodactyla, Primates, and Proboscidea) and discuss their paleoecological and biochronological significance. The size of the Maka'amitalu assemblage is small compared to those from the Hadar Formation (3.45-2.95 Ma) and Ledi-Geraru (2.8-2.6 Ma) but includes at least 20 taxa. Bovids, suids, and Theropithecus are common in terms of both species richness and abundance, whereas carnivorans, equids, and megaherbivores are rare. While the taxonomic composition of the Maka'amitalu fauna indicates significant species turnover from the Hadar Formation and Ledi-Geraru deposits, turnover seems to have occurred at a constant rate through time as taxonomic dissimilarity between adjacent fossil assemblages is strongly predicted by their age difference. A similar pattern characterizes functional ecological turnover, with only subtle changes in dietary proportions, body size proportions, and bovid abundances across the composite lower Awash sequence. Biochronological comparisons with other sites in eastern Africa suggest that the taxa recovered from the Maka'amitalu are broadly consistent with the reported age of the BKT-3 tuff. Considering the age of BKT-3 and biochronology, a range of 2.4-1.9 Ma is most likely for the faunal assemblage.

Plio-Pleistocene environmental variability in Africa and its implications for mammalian evolution.

Understanding the climatic drivers of environmental variability (EV) during the Plio-Pleistocene and EV's influence on mammalian macroevolution are two outstanding foci of research in African paleoclimatology and evolutionary biology. The potential effects of EV are especially relevant for testing the variability selection hypothesis, which predicts a positive relationship between EV and speciation and extinction rates in fossil mammals. Addressing these questions is stymied, however, by 1) a lack of multiple comparable EV records of sufficient temporal resolution and duration, and 2) the incompleteness of the mammalian fossil record. Here, we first compile a composite history of Pan-African EV spanning the Plio-Pleistocene, which allows us to explore which climatic variables influenced EV. We find that EV exhibits 1) a long-term trend of increasing variability since ∼3.7 Ma, coincident with rising variability in global ice volume and sea surface temperatures around Africa, and 2) a 400-ky frequency correlated with seasonal insolation variability. We then estimate speciation and extinction rates for fossil mammals from eastern Africa using a method that accounts for sampling variation. We find no statistically significant relationship between EV and estimated speciation or extinction rates across multiple spatial scales. These findings are inconsistent with the variability selection hypothesis as applied to macroevolutionary processes.

No sustained increase in zooarchaeological evidence for carnivory after the appearance of Homo erectus.

The appearance of Homo erectus shortly after 2.0 Ma is widely considered a turning point in human dietary evolution, with increased consumption of animal tissues driving the evolution of larger brain and body size and a reorganization of the gut. An increase in the size and number of zooarchaeological assemblages after the appearance of H. erectus is often offered as a central piece of archaeological evidence for increased carnivory in this species, but this characterization has yet to be subject to detailed scrutiny. Any widespread dietary shift leading to the acquisition of key traits in H. erectus should be persistent in the zooarchaeological record through time and can only be convincingly demonstrated by a broad-scale analysis that transcends individual sites or localities. Here, we present a quantitative synthesis of the zooarchaeological record of eastern Africa from 2.6 to 1.2 Ma. We show that several proxies for the prevalence of hominin carnivory are all strongly related to how well the fossil record has been sampled, which constrains the zooarchaeological visibility of hominin carnivory. When correcting for sampling effort, there is no sustained increase in the amount of evidence for hominin carnivory between 2.6 and 1.2 Ma. Our observations undercut evolutionary narratives linking anatomical and behavioral traits to increased meat consumption in H. erectus, suggesting that other factors are likely responsible for the appearance of its human-like traits.

Rethinking the ecological drivers of hominin evolution.

A central goal of paleoanthropology is understanding the role of ecological change in hominin evolution. Over the past several decades researchers have expanded the hominin fossil record and assembled detailed late Cenozoic paleoclimatic, paleoenvironmental, and paleoecological archives. However, effective use of these data is precluded by the limitations of pattern-matching strategies for inferring causal relationships between ecological and evolutionary change. We examine several obstacles that have hindered progress, and highlight recent research that is addressing them by (i) confronting an incomplete fossil record, (ii) contending with datasets spanning varied spatiotemporal scales, and (iii) using theoretical frameworks to build stronger inferences. Expanding on this work promises to transform challenges into opportunities and set the stage for a new phase of paleoanthropological research.

Intrataxonomic trends in herbivore enamel δ13C are decoupled from ecosystem woody cover.

Analysis of enamel stable carbon isotopes (δ13Cenamel) of fossil herbivores is an important tool for making inferences about Plio-Pleistocene vegetation structure in Africa and the environmental context of hominin evolution. Many palaeoecological studies implicitly or explicitly assume that individual variation in C3-C4 plant consumption among fossil herbivores directly reflects the abundance of C3 (trees, shrubs) or C4 (low-altitude tropical grasses) vegetation. However, a strong link between δ13Cenamel of herbivores and ecosystem vegetation structure has not been rigorously established. Here we combine δ13Cenamel data from a large dataset (n = 1,643) with multidecadal Landsat estimates of C3 woody cover across 30 African ecosystems to show that there is little relationship between intrataxonomic variation in δ13Cenamel and vegetation structure. This is especially true when removing forested ecosystems (>80% woody cover)-which numerous lines of evidence suggest are rare in the Plio-Pleistocene fossil record of eastern Africa-from our analyses. Our findings stand in contrast with the common assumption that variation in herbivore δ13Cenamel values reflects changes in the relative abundance of C3-C4 vegetation. We conclude that analyses using herbivore δ13Cenamel data to shed light on the environmental context of hominin evolution should look to explicitly community-level approaches for making vegetation inferences.

Functional traits of the world's late Quaternary large-bodied avian and mammalian herbivores.

Prehistoric and recent extinctions of large-bodied terrestrial herbivores had significant and lasting impacts on Earth's ecosystems due to the loss of their distinct trait combinations. The world's surviving large-bodied avian and mammalian herbivores remain among the most threatened taxa. As such, a greater understanding of the ecological impacts of large herbivore losses is increasingly important. However, comprehensive and ecologically-relevant trait datasets for extinct and extant herbivores are lacking. Here, we present HerbiTraits, a comprehensive functional trait dataset for all late Quaternary terrestrial avian and mammalian herbivores ≥10 kg (545 species). HerbiTraits includes key traits that influence how herbivores interact with ecosystems, namely body mass, diet, fermentation type, habitat use, and limb morphology. Trait data were compiled from 557 sources and comprise the best available knowledge on late Quaternary large-bodied herbivores. HerbiTraits provides a tool for the analysis of herbivore functional diversity both past and present and its effects on Earth's ecosystems.

Evaluation of chromosomal insertion loci in the Pseudomonas putida KT2440 genome for predictable biosystems design.

The development of Pseudomonas strains for industrial production of fuels and chemicals will require the integration of heterologous genes and pathways into the chromosome. Finding the most appropriate integration site to maximize strain performance is an essential part of the strain design process. We characterized seven chromosomal loci in Pseudomonas putida KT2440 for integration of a fluorescent protein expression construct. Insertion in five of the loci did not affect growth rate, but fluorescence varied by up to 27-fold. Three sites displaying a diversity of phenotypes with the fluorescent reporter were also chosen for the integration of a gene encoding a muconate importer. Depending on the integration locus, expression of the importer varied by approximately 3-fold and produced significant phenotypic differences. This work demonstrates the impact of the integration location on host viability, gene expression, and overall strain performance.

Introduced herbivores restore Late Pleistocene ecological functions.

Large-bodied mammalian herbivores dominated Earth's terrestrial ecosystems for several million years before undergoing substantial extinctions and declines during the Late Pleistocene (LP) due to prehistoric human impacts. The decline of large herbivores led to widespread ecological changes due to the loss of their ecological functions, as driven by their unique combinations of traits. However, recently, humans have significantly increased herbivore species richness through introductions in many parts of the world, potentially counteracting LP losses. Here, we assessed the extent to which introduced herbivore species restore lost-or contribute novel-functions relative to preextinction LP assemblages. We constructed multidimensional trait spaces using a trait database for all extant and extinct mammalian herbivores ≥10 kg known from the earliest LP (∼130,000 ybp) to the present day. Extinction-driven contractions of LP trait space have been offset through introductions by ∼39% globally. Analysis of trait space overlap reveals that assemblages with introduced species are overall more similar to those of the LP than native-only assemblages. This is because 64% of introduced species are more similar to extinct rather than extant species within their respective continents. Many introduced herbivores restore trait combinations that have the capacity to influence ecosystem processes, such as wildfire and shrub expansion in drylands. Although introduced species have long been a source of contention, our findings indicate that they may, in part, restore ecological functions reflective of the past several million years before widespread human-driven extinctions.

Trophic rewilding presents regionally specific opportunities for mitigating climate change.

Large-bodied mammalian herbivores can influence processes that exacerbate or mitigate climate change. Herbivore impacts are, in turn, influenced by predators that place top-down forcing on prey species within a given body size range. Here, we explore how the functional composition of terrestrial large-herbivore and -carnivore guilds varies between three mammal distribution scenarios: Present-Natural, Current-Day and Extant-Native Trophic (ENT) Rewilding. Considering the effects of herbivore species weakly influenced by top-down forcing, we quantify the relative influence keystone large-herbivore guilds have on methane emissions, woody vegetation expansion, fire dynamics, large-seed dispersal, and nitrogen and phosphorus transport potential. We find strong regional differences in the number of herbivores under weak top-down regulation between our three scenarios, with important implications for how they will influence climate change relevant processes. Under the Present-Natural non-ruminant, megaherbivore, browsers were a particularly important guild across much of the world. Megaherbivore extinction and range contraction and the arrival of livestock mean large, ruminant, grazers have become more dominant. ENT Rewilding can restore the Afrotropics and the Indo-Malay realm to the Present-Natural benchmark, but causes top-down forcing of the largest herbivores to become commonplace elsewhere. ENT Rewilding will reduce methane emissions, but does not maximize natural climate solution potential. This article is part of the theme issue 'Climate change and ecosystems: threats, opportunities and solutions'.

Statistical estimates of hominin origination and extinction dates: A case study examining the Australopithecus anamensis-afarensis lineage.

Reliable estimates of when hominin taxa originated and went extinct are central to addressing many paleoanthropological questions, including those relating to macroevolutionary patterns. The timing of hominin temporal ranges can be used to test chronological predictions generated from phylogenetic hypotheses. For example, hypotheses of phyletic ancestor-descendant relationships, based on morphological data, predict no temporal range overlap between the two taxa. However, a fossil taxon's observed temporal range is almost certainly underestimated due to the incompleteness of both the fossil record itself and its sampling, and this decreases the likelihood of observing temporal overlap. Here, we focus on a well-known and widely accepted early hominin lineage, Australopithecus anamensis-afarensis, and place 95% confidence intervals (CIs) on its origination and extinction dates. We do so to assess whether its temporal range is consistent with it being a phyletic descendant of Ardipithecus ramidus and/or a direct ancestor to the earliest claimed representative of Homo (i.e., Ledi-Geraru). We find that the last appearance of Ar. ramidus falls within the origination CI of Au. anamensis-afarensis, whereas the claimed first appearance of Homo postdates the extinction CI. These results are consistent with Homo evolving from Au. anamensis-afarensis, but temporal overlap between Ar. ramidus and Au. anamensis-afarensis cannot be rejected at this time. Though additional samples are needed, future research should extend our initial analyses to incorporate the uncertainties surrounding the range endpoints of Ar. ramidus and earliest Homo. Overall, our findings demonstrate the need for quantifying the uncertainty surrounding the appearances and disappearances of hominin taxa in order to better understand the timing of evolutionary events in our clade's history.

Geographically divergent evolutionary and ecological legacies shape mammal biodiversity in the global tropics and subtropics.

Studies of the factors governing global patterns of biodiversity are key to predicting community responses to ongoing and future abiotic and biotic changes. Although most research has focused on present-day climate, a growing body of evidence indicates that modern ecological communities may be significantly shaped by paleoclimatic change and past anthropogenic factors. However, the generality of this pattern is unknown, as global analyses are lacking. Here we quantify the phylogenetic and functional trait structure of 515 tropical and subtropical large mammal communities and predict their structure from past and present climatic and anthropogenic factors. We find that the effects of Quaternary paleoclimatic change are strongest in the Afrotropics, with communities in the Indomalayan realm showing mixed effects of modern climate and paleoclimate. Malagasy communities are poorly predicted by any single factor, likely due to the atypical history of the island compared with continental regions. Neotropical communities are mainly codetermined by modern climate and prehistoric and historical human impacts. Overall, our results indicate that the factors governing tropical and subtropical mammalian biodiversity are complex, with the importance of past and present factors varying based on the divergent histories of the world's biogeographic realms and their native biotas. Consideration of the evolutionary and ecological legacies of both the recent and ancient past are key to understanding the forces shaping global patterns of present-day biodiversity and its response to ongoing and future abiotic and biotic changes in the 21st century.

Early hominins evolved within non-analog ecosystems.

Present-day African ecosystems serve as referential models for conceptualizing the environmental context of early hominin evolution, but the degree to which modern ecosystems are representative of those in the past is unclear. A growing body of evidence from eastern Africa's rich and well-dated late Cenozoic fossil record documents communities of large-bodied mammalian herbivores with ecological structures differing dramatically from those of the present day, implying that modern communities may not be suitable analogs for the ancient ecosystems of hominin evolution. To determine when and why the ecological structure of eastern Africa's herbivore faunas came to resemble those of the present, here we analyze functional trait changes in a comprehensive dataset of 305 modern and fossil herbivore communities spanning the last ∼7 Myr. We show that nearly all communities prior to ∼700 ka were functionally non-analog, largely due to a greater richness of non-ruminants and megaherbivores (species >1,000 kg). The emergence of functionally modern communities precedes that of taxonomically modern communities by 100,000s of years, and can be attributed to the combined influence of Plio-Pleistocene C4 grassland expansion and pulses of aridity after ∼1 Ma. Given the disproportionate ecological impacts of large-bodied herbivores on factors such as vegetation structure, hydrology, and fire regimes, it follows that the vast majority of early hominin evolution transpired in the context of ecosystems that functioned unlike any today. Identifying how past ecosystems differed compositionally and functionally from those today is key to conceptualizing ancient African environments and testing ecological hypotheses of hominin evolution.