Large Mammalian Herbivores Research Articles

Comparative study of the body proportions in Elephantidae and other large herbivorous mammals.

In this study, we aimed to achieve three objectives: (1) to precisely characterize the body plans ofElephantidae and other large herbivorous mammals; (2) based on this analysis, to determine whether the body plans of the extinct woolly mammoth (Mammuthus primigenius) and steppe mammoth (M. trogontherii) differ from those of modern-day Elephantidae: the Asian elephant (Elephas maximus), the African bush (Loxodonta africana), and forest (L. cyclotis) elephants; (3) to analyze how the body plans have changed in extant perissodactyls and proboscideans compared with their Paleogene ancestors. To accomplish this, we studied mammoth skeletons from the collections of Russian museums and compared this data with a large number of skeletons of extant elephantids, odd-toed, and even-toed ungulates, as well as their extinct relatives. We showed that three genera of Elephantidae are characterized by a homogeneous body plan, which is markedly different from other large herbivores. Elephantids break the interrelationship, that exists in artiodactyls and perissodactyls, between the total length of the head and neck on one side and the limb's segments on the other. Their limbs are very tall (inferior in this regard among large ungulates only to the giraffe), and, contrary to the other large herbivorous mammals, elongated due to the length of the proximal segments. This allows them to effectively utilize the principle of inverted pendulum (straight-legged walking) in locomotion. The biggest differences in the body plan of mammoths compared with extant elephants are a markedly larger pelvis, elongated fore- and hindlimbs (due to the increased relative length of their proximal segments), and different proportions of the skull. The body plans of plesiomorphic Paleogene proboscideans and perissodactyls differed markedly from their descendants in every body part; these differences are related, on the one hand, to the allometric growth, and on the other hand, to the advancement of the locomotor apparatus in the course of their evolution. The most notable difference in the body plan between Paleogene proboscidean Moeritherium and extant Elephantidae is the ~2-fold increase in relative limb height.

Behavioural responses to mammalian grazing expose insect herbivores to elevated risk of avian predation.

Large mammalian herbivores (LMH) are important functional components and drivers of biodiversity and ecosystem functioning in grasslands. Yet their role in regulating food-web dynamics and trophic cascades remains poorly understood. In the temperate grasslands of northern China, we explored whether and how grazing domestic cattle (Bos taurus) alter the predator-prey interactions between a dominant grasshopper (Euchorthippus unicolor) and its avian predator the barn swallow (Hirundo rustica). Using two large manipulative field experiments, we found that in the presence of cattle, grasshoppers increased their jumping frequency threefold, swallows increased foraging visits to these fields sixfold, and grasshopper density was reduced by about 50%. By manipulatively controlling the grasshoppers' ability to jump, we showed that jumping enables grasshoppers to avoid being incidentally consumed or trampled by cattle. However, jumping behaviour increased their consumption rates by swallows 37-fold compared with grasshoppers that were unable to jump. Our findings illustrate how LMH can indirectly alter predator-prey interactions by affecting behaviour of avian predators and herbivorous insects. These non-plant-mediated effects of LMH may influence trophic interactions in other grazing ecosystems and shape community structure and dynamics. We highlight that convoluted multispecies interactions may better explain how LMH control food-web dynamics in grasslands.

Higher abundance of disturbance‐favoured trees and shrubs in European temperate woodlands prior to the late‐quaternary extinction of megafauna

Abstract Large herbivorous mammals strongly influence vegetation structure by creating and maintaining open areas and causing disturbance within closed woody habitats. The herbivores alive today in Europe are only a small remnant of the large species that existed in high diversity and abundance before modern humans. The extinction of so many large herbivores during the last 50,000 years, and the loss of megaherbivores (body weight ≥1000 kg) from most of the continent before the Pleistocene–Holocene transition, is likely to have had cascading effects on vegetation structure and composition. To evaluate these effects within the European temperate forest biome, we examine the abundance change of three important European woody taxa (deciduous oaks, Quercus spp.; hazel, Corylus [mainly C. avellana]; and yew, Taxus baccata) before and after the late‐Quaternary downgrading of the region''s large herbivore fauna. These taxa are disturbance‐favoured, depend on canopy openings for regeneration, and tend to decline in closed dense forests. Quercus and Corylus may thrive in systems affected by megafauna herbivory or fire, whilst Taxus is fire‐sensitive but can thrive in grazed systems. Using pollen‐based reconstructions (REVEALS), we investigated the proportional cover of these three focal taxa in the Last Interglacial (129,000–116,000 before present [BP]) and early–mid‐Holocene (8700–5700 BP). We found that woodlands in the Last Interglacial exhibited higher cover of Corylus and Taxus relative to the Holocene, with the former reaching very high percentage cover; meanwhile, Quercus had a consistent, moderately high percentage cover in both periods. Furthermore, we found that the cover of Corylus and Taxus appeared to be influenced more by unmeasured, non‐climatic factors than Quercus. Synthesis: The abundance of Taxus suggests a limited role of fire; whilst the observed levels of Taxus, Corylus and Quercus align with the potential influence of megafauna herbivory; however, a direct quantitative link remains to be established. Our results suggest that vegetation was structurally and compositionally affected by differences in disturbance regimes between the Last Interglacial and early–mid‐Holocene, with the loss of diverse disturbance regimes likely contributing to the divergence of Holocene vegetation from long‐term ecological baselines.

Body size modulates the extent of seasonal diet switching by large mammalian herbivores in Yellowstone National Park.

Prevailing theories about animal foraging behaviours and the food webs they occupy offer divergent predictions about whether seasonally limited food availability promotes dietary diversification or specialization. Emphasis on how animals compete for food predominates in work on the foraging ecology of large mammalian herbivores, whereas emphasis on how the diversity of available foods generally constrains dietary opportunity predominates work on entire food webs. Reconciling predictions about what promotes dietary diversification is challenging because species' different body sizes and mobilities modulate how they seek and compete for resources-the mechanistic bases of common predictions may not pertain to all species equally. We evaluated predictions about five large-herbivore species that differ in body size and mobility in Yellowstone National Park using GPS tracking and dietary DNA. The data illuminated remarkably strong and significant correlations between body size and five key indicators of diet seasonality (R 2 = 0.71-0.80). Compared to smaller species, bison and elk showed muted diet seasonality and maintained access to more unique foods when winter conditions constrained food availability. Evidence from GPS collars revealed size-based differences in species' seasonal movements and habitat-use patterns, suggesting that better accounting for the allometry of foraging behaviours may help reconcile disparate ideas about the ecological drivers of seasonal diet switching.

Baring Bearma’s Bounty. Human and animal signatures in the Quaternary contexts of the Bearma Valley, Central India

The rich Palaeolithic and fossil records of the Narmada and Son Valleys dominate paleoanthropological research from Central India, in large part because of their well-preserved Quaternary deposits. While data from other river valleys is emerging, finding both lithics and fauna in close proximity remains rare and spatio-temporal inconsistencies make assessment of implications for the larger area challenging. Ongoing investigations along the Bearma, the largest tributary of the River Sonar, has yielded a low-density fossil assemblage across three localities and a proximally located Large Flake Acheulean (LFA) site. The small fossil assemblage currently consists of both cranial and post-cranial elements in differing states of completeness and preservation, likely a function of post-depositional impacts and the time elapsed since exposure. Many of the important large mammalian herbivore species belonging to bovid, equid, cervid and proboscidean taxa that are commonly found during the Late Quaternary of the Indian Subcontinent are represented, making the region worthy of comparison with neighbouring fossiliferous areas. The new Palaeolithic site encountered less than 500 m downstream from the fossil localities is almost exclusively LFA technology in nature made primarily on slabs of Vindhyan sandstone. Preliminary analysis of the collected artefacts indicates a focus on the production of large flake blanks and debitage products from large cores configured according to various unifacial and bifacial technological methods. The frequency of these elements suggests the use of the area as a factory site, while the unmistakable presence of bifaces and other tools warrant broader interpretations. With the current lack of stratigraphic and chronometric data on either assemblage, it is premature to draw any chrono-contextual associations between the fossils and artefacts. However, this ∼5 km stretch of the Bearma yielding two significant proxies presents an excellent opportunity to understand hominin behaviour during the Quaternary and fill both geographical and temporal gaps in Central India.

Tree growth‐forms reveal dominant browsers shaping the vegetation

Abstract Plants adopt particular growth‐forms when they are exposed to extreme environmental conditions. In this study, we describe a unique woody plant growth‐form induced by large mammalian herbivores and discuss that this growth‐form could have evolved as a strategy for escaping the browser zone in herbivore driven ecosystems. We analysed responses of key architectural and morphological attributes (branching and thorn density, tree dimensions, presence of flowers and fruits) of three Eurasian spiny tree species (Malus sylvestris, Prunus cerasifera and Pyrus communis) to different levels of browsing by large herbivores in the temperate Białowieża Forest, Poland. Under high browsing pressure, studied trees displayed two distinct forms of the crown: a bottom sterile part developing into a densely branched structure with high density of thorns (‘cage‐form’), and an upper reproductive part that escaped from herbivore control (‘escaped‐form’). The size of cage‐form influenced the feeding behaviour of red deer (Cervus elaphus) by increasing the time deer spend foraging and increasing the bite rate. The height at which cages started to escape and their diameter matched with foraging reach of red deer. Synthesis. We argue that the frequency and cage dimensions of this woody growth‐form in the landscape could inform on the type and intensity of recent herbivory. Moreover, its distinctive inducibility suggests that this growth‐form did not emerge recently under anthropogenic pressure but could be the legacy of ancient herbivory effects. Observational evidence suggests that this growth‐form emerged in several herbivore‐driven systems around the globe and may be used to identify the dominant herbivores that control vegetation structure in these ecosystems. Read the free Plain Language Summary for this article on the Journal blog.

Environmental DNA metabarcoding reveals temporal dynamics but functional stability of arthropod communities in cattle dung.

Terrestrial invertebrates are highly important for the decomposition of dung from large mammals. Mammal dung has been present in many of Earth's ecosystems for millions of years, enabling the evolution of a broad diversity of dung-associated invertebrates that process various components of the dung. Today, large herbivorous mammals are increasingly introduced to ecosystems with the aim of restoring the ecological functions formerly provided by their extinct counterparts. However, we still know little about the ecosystem functions and nutrient flows in these rewilded ecosystems, including the dynamics of dung decomposition. In fact, the succession of insect communities in dung is an area of limited research attention also outside a rewilding context. In this study, we use environmental DNA metabarcoding of dung from rewilded Galloway cattle in an experimental set-up to investigate invertebrate communities and functional dynamics over a time span of 53 days, starting from the time of deposition. We find a strong signal of successional change in community composition, including for the species that are directly dependent on dung as a resource. While several of these species were detected consistently across the sampling period, others appeared confined to either early or late successional stages. We believe that this is indicative of evolutionary adaptation to a highly dynamic resource, with species showing niche partitioning on a temporal scale. However, our results show consistently high species diversity within the functional groups that are directly dependent on dung. Our findings of such redundancy suggest functional stability of the dung-associated invertebrate community, with several species ready to fill vacant niches if other species disappear. Importantly, this might also buffer the ecosystem functions related to dung decomposition against environmental change. Interestingly, alpha diversity peaked after approximately 20-25 days in both meadow and pasture habitats, and did not decrease substantially during the experimental period, probably due to preservation of eDNA in the dung after the disappearance of visiting invertebrates, and from detection of tissue remains and cryptic life stages.

Ecosystem‐wide responses to fire and large mammal herbivores in an African savanna

AbstractFire and large mammal herbivores (LMH) are the principal top‐down forces maintaining savanna structure. Nonetheless, experiments designed to investigate interactions between fire and LMH are rare in savannas, and relationships between environmental variation and biodiversity in the context of fire and LMH are poorly understood. This study addresses these gaps by manipulating the presence of LMH and early and late dry season fires in a tropical African savanna. In addition, this work simultaneously explores environmental variables including soil and foliar quality, vegetation cover, and nearby water sources to more holistically describe factors affecting savanna functioning and biodiversity. After 1 year of experimental treatments, changes in vegetation were already apparent. Shrub abundance and richness and grass richness were higher in the absence of LMH, while grass biomass increased three‐fold in burned plots as compared to unburned plots. Foliar nutrients tended to increase in open plots, while phenolics decreased. Amphibian abundance decreased with early burns and was higher with LMH. In contrast, small mammal abundance and richness increased without LMH and with time since fire. Richness and foraging of LMH were highest after late burns. These results demonstrate ecosystem‐wide effects of LMH, illustrating the importance of considering multiple taxa when designing fire management programs. For example, burning negatively affected amphibians and small mammals and changed vegetation at the same time it increased LMH foraging. In the long‐term, this experiment will shed light on interacting effects of fire and LMH on savanna biodiversity and function.Abstract in Portuguese is available with the online material.

A theory for context-dependent effects of mammalian trampling on ecosystem nitrogen cycling.

Large mammalian herbivores substantially impact ecosystem functioning. As their populations are dramatically altered globally, disentangling their consumptive and non-consumptive effects is critical to advance mechanistic understanding and improve prediction of effects over ecosystem and Earth-system spatial extents. Mathematical models have played an important role in clarifying potential mechanisms of herbivore zoogeochemistry, based mostly on their consumptive effects as primary consumers and recyclers of organic and inorganic matter via defecation and urination. Trampling is a ubiquitous effect among walking vertebrates, but the consequences and potential mechanisms of trampling in diverse environments remain poorly understood. We derive a novel mathematical model of large mammalian herbivore effects on ecosystem nitrogen cycling, focusing on how trampling and environmental context impact soil processes. We model herbivore trampling with a linear positive or negative additive effect on soil-mediated nitrogen cycling processes. Combining analytical and numerical analyses, we find trampling by large mammalian herbivores is likely to decrease nitrogen mineralisation rate across diverse environments, such as temperate grassland and boreal forest. These effects are mediated by multiple potential mechanisms, including trampling-induced changes to detritivore biomass and functioning (e.g. rate of organic matter consumption). We also uncover scenarios where trampling can increase nitrogen mineralisation rate, contingent on the environment-specific relative sensitivity of detritivore mineral-nitrogen release and detritivore mortality, to trampling. In contrast to some consumptive mechanisms, our results suggest the pace of soil nitrogen cycling prior to trampling has little influence over the direction of the trampling net effect on nitrogen mineralisation, but that net effects may be greater in slow-cycling systems (e.g. boreal forests) than in fast-cycling systems (e.g. grasslands). Our model clarifies the potential consequences of previously overlooked mechanisms of zoogeochemistry that are common to all terrestrial biomes. Our results provide empirically testable predictions to guide future progress in empirical and theoretical studies of herbivore effects in diverse environmental contexts. Resolving ecological contingencies around animal consumptive and non-consumptive effects will improve whole-ecosystem management efforts such as restoration and rewilding.

Blocking then stinging as a case of two-step evolution of defensive cage architectures in herbivore-driven ecosystems.

Dense branching and spines are common features of plant species in ecosystems with high mammalian herbivory pressure. While dense branching and spines can inhibit herbivory independently, when combined, they form a powerful defensive cage architecture. However, how cage architecture evolved under mammalian pressure has remained unexplored. Here we show how dense branching and spines emerged during the age of mammalian radiation in the Combretaceae family and diversified in herbivore-driven ecosystems in the tropics. Phylogenetic comparative methods revealed that modern plant architectural strategies defending against large mammals evolved via a stepwise process. First, dense branching emerged under intermediate herbivory pressure, followed by the acquisition of spines that supported higher speciation rates under high herbivory pressure. Our study highlights the adaptive value of dense branching as part of a herbivore defence strategy and identifies large mammal herbivory as a major selective force shaping the whole plant architecture of woody plants.

The interplay between defaunation and phylogenetic diversity affects leaf damage by natural enemies in tropical plants

Abstract Natural enemies play an important role in controlling plant population growth and vegetation dynamics. Tropical rainforests host the greatest diversity of herbivores, from large mammalian ungulates to microscopic pathogens, generating and maintaining plant diversity. By feeding on the same resources, large mammalian herbivores may interfere with plant consumption and leaf damage by important enemy guilds such as invertebrate herbivores and pathogens, triggering indirect trophic cascades. However, the impact of local extinctions of large herbivores on plant–enemy interactions is relatively unknown. We experimentally tested the effects of defaunation of large mammalian herbivores (e.g. peccaries, tapirs and brocket deer; hereafter, large herbivores) on the leaf damage of 3350 understorey plants in tropical rainforests of Brazil. We examined leaf damage in 10,050 leaves from 333 morphospecies by assigning the area consumed or damaged by five guilds of insect herbivores and leaf pathogens within 86 paired open‐closed plots and investigating the joint effects of defaunation and plant phylogenetic diversity. Plants released from large herbivores had 9% less leaf damage; this difference was due to the lower leaf pathogens incidence (29%) rather than insect herbivory. Evolutionary distinctness was positively correlated with leaf damage in a similar way in all treatments, suggesting additive effects of defaunation and phylogenetic diversity. Total and pathogenic leaf damage (but not insect damage) decreased with plant richness across treatments, and large herbivores exclusion resulted in increased plant species richness. This suggests that large herbivores exclusion leads to a dilution of total and pathogens' leaf damage by increasing plant species richness. Our results suggest that indirect effects of large herbivores decrease the dilution potential of plant communities against pathogens and rather reinforce their top‐down impact on vegetation, demonstrating a previously overlooked cascading effect of large herbivore extinction on forest ecosystems. Synthesis: The extinction of large mammalian herbivores can lead to a decrease in pathogen‐driven leaf damage, a previously unknown indirect effect in forest ecosystems, which might have consequences for plant fitness and ultimately for plant diversity. Large herbivores and plant pathogens might have synergistic effects in regulating the diversity of plant communities in some of the most diverse ecosystems on Earth.

Functional traits-not nativeness-shape the effects of large mammalian herbivores on plant communities.

Large mammalian herbivores (megafauna) have experienced extinctions and declines since prehistory. Introduced megafauna have partly counteracted these losses yet are thought to have unusually negative effects on plants compared with native megafauna. Using a meta-analysis of 3995 plot-scale plant abundance and diversity responses from 221 studies, we found no evidence that megafauna impacts were shaped by nativeness, "invasiveness," "feralness," coevolutionary history, or functional and phylogenetic novelty. Nor was there evidence that introduced megafauna facilitate introduced plants more than native megafauna. Instead, we found strong evidence that functional traits shaped megafauna impacts, with larger-bodied and bulk-feeding megafauna promoting plant diversity. Our work suggests that trait-based ecology provides better insight into interactions between megafauna and plants than do concepts of nativeness.

The role of large mammalian herbivores in shaping and maintaining soil microbial communities of natural mineral licks: A case study on sika deer at the firebreak adjacent to the Sino-Russian border.

Mineral licks are indispensable habitats to the life history of large mammal herbivores (LMH). Geophagy at licks may provide the necessary minerals for LMH, while LMH may be ecosystem engineers of licks by altering vegetation cover and soil physicochemical properties (SPCP). However, the precise relationship between the LMH and licks remains unclear. To clarify the geophagy function of licks for LMH and their influence on soil at licks, we recorded visitation patterns of sika deer around licks and compared SPCP and microbial communities with the surrounding matrix in a firebreak adjacent to the Sino-Russian border. Our study indirectly supports the "sodium supplementation" hypothesis. Proofs included (1) a significantly higher sodium, iron, and aluminum contents than the matrix, while lower carbon, nitrogen, and moisture contents; (2) significantly higher deer visitation during sodium-demand season (growing season), along with an avoidance of licks with high iron contents, which is toxic when overdose. The microbes at the licks differed from those at the matrix, mainly driven by low soil carbon and nitrogen and altered biogeochemical cycles. The microbial communities of licks are vulnerable because of their unstable state and susceptibility to SPCP changes. Structural equation modeling (SEM) clearly showed a much stronger indirect effect of deer on microbes at licks than at the matrix, especially for bacteria. Multiple deer behaviors at licks, such as grazing, trampling, and excretion, can indirectly shape and stabilize microbes by altering carbon and nitrogen input. Our study is the first to characterize soil microbial communities at mineral licks and demonstrate the processes by which LMH shapes those communities. More studies are required to establish a general relationship between the LMH and licks to promote the conservation of natural licks for wildlife.

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.

Uncovering widespread Anthropocene dietary shifts in Chinese large mammalian herbivores.

The Anthropocene's human-dominated habitat expansion endangers global biodiversity. However, large mammalian herbivores experienced few extinctions during the 20th century, hinting at potentially overlooked ecological responses of a group sensitive to global change. Using dental microwear as a proxy, we studied large herbivore dietary niches over a century across mainland China before (1880s-1910s) and after (1970s-1990s) the human population explosion. We uncovered widespread and significant shifts (interspecific microwear differences increased and intraspecific microwear dispersion expanded) within dietary niches linked to geographical areas with rapid industrialization and population growth in eastern China. By contrast, in western China, where human population growth was slower, we found no indications of shifts in herbivore dietary niches. Further regression analysis links the intensity of microwear changes to human land-use expansion. These analyses highlight dietary adjustments of large herbivores as a likely key factor in their adaptation across a century of large-scale human-driven changes.

Paleobiology of Pleistocene large land mammals from the Brazilian Pampa

The southern Brazilian Pleistocene fauna exhibited a rich diversity of large mammalian herbivores, which are now extinct or locally extinct. In this study, we employed stable isotope analysis to investigate the past ecology of these animals. Specifically, we examined the carbonate fraction of bones and teeth and utilized compiled carbon and oxygen isotopic ratios from previous research publications. The δ13C values indicated that most specimens inhabited grassland environments, which aligns with environmental reconstructions based on pollen records of the “Campos” region. The dominant food resource for these herbivores consisted of C3 photosynthesizers, mainly cool-season grasses. This preference can be attributed to the higher abundance and nutritional quality of cool-season grasses compared with warm-season grasses employing C4 photosynthesis. The variability in δ18O values within and between taxa may suggest a seasonal climate. Based on these findings, we conclude that the environmental changes during the Pleistocene-Holocene transition were detrimental to the survival of these large herbivores.

Community composition, and not species richness, of microbes influences decomposer functional diversity in soil

Diversity-function relationships in producers, and how these are influenced by consumers, are well known. However, these are not well known for microbial decomposers in soil. It is also unknown whether and how consumers such as large mammalian herbivores influence soil microbial decomposer diversity-function relationships. We used a 14-year-old herbivore-exclusion experiment in the Trans-Himalayan drylands of northern India to address whether microbial functions vary with microbial diversity (both species richness and composition), and whether herbivores alter the diversity-function relationships. We analyzed soils from n = 10 paired grazed-and-fenced plots three times during the growth season of 2019. Data were from 16S rDNA gene amplicon sequencing of 7.6 million reads covering 1937 operational taxonomic units (OTU) of bacteria across 47 phyla (of which 924 OTUs were identifiable to genus level), and 1800 catabolic profiles across 30 substrates related to carbon metabolism. We found that functional diversity was positively related to microbial community composition, but not to species richness; it was also unaffected by large mammalian herbivore-exclusion to indicate resilience and resistance. This positive relationship between community composition and functional diversity challenges the prevailing notion of functional redundancy in hyper-diverse soil microbial communities since certain combinations of species could outperform others and determine decomposition processes and services. Structural equation models suggested that the strength of this relationship is favoured by availability of soil moisture. Microbial functions varied more strongly with temporal variables (e.g., seasonality) than with spatial variables (e.g., edaphic factors such as soil texture and pH). Although interpretations can be constrained by which and how many functions are investigated, the relationship was generalizable and robust once 16 or more functions were quantified. Decomposition in drylands may be particularly susceptible to how the identity of the microbial species, and not the number of species, responds to rising precipitation variability under ongoing and projected climate change.

Paleoenvironmental inferences on the Late Miocene hominoid-bearing site of Can Llobateres (NE Iberian Peninsula): An ecometric approach based on functional dental traits

Hispanopithecus laietanus from the Late Miocene (9.8 Ma) of Can Llobateres 1 (CLL1; Vallès-Penedès Basin, NE Iberian Peninsula) represents one of the latest occurrences of fossil apes in Western mainland Europe, where they are last recorded at ∼9.5 Ma. The paleoenvironment of CLL1 is thus relevant for understanding the extinction of European hominoids. To refine paleoenvironmental inferences for CLL1, we apply ecometric models based on functional crown type (FCT) variables—a scoring scheme devised to capture macroscopic functional traits of occlusal shape and wear surfaces of herbivorous large mammal molars. Paleotemperature and paleoprecipitation estimates for CLL1 are provided based on published regional regression models linking average FCT of large herbivorous mammal communities to climatic conditions. A mapping to Whittaker's present-day biome classification is also attempted based on these estimates, as well as a case-based reasoning via canonical variate analysis of FCT variables from five relevant biomes. Estimates of mean annual temperature (25 °C) and mean annual precipitation (881 mm) classify CLL1 as a tropical seasonal forest/savanna, only in partial agreement with the canonical variate analysis results, which classify CLL1 as a tropical rainforest with a higher probability. The former biome agrees better with previous inferences derived from fossil plants and mammals, as well as preliminary isotopic data. The misclassification of CLL1 as a tropical forest is attributed to the mixture of forest-adapted taxa with others adapted to more open environments, given that faunal and plant composition indicates the presence of a dense wetland/riparian forest with more open woodlands nearby. The tested FCT ecometric approaches do not provide unambiguous biome classification for CLL1. Nevertheless, our results are consistent with those from other approaches, thus suggesting that FCT variables are potentially useful to investigate paleoenvironmental changes through time and space—including those that led to the extinction of European Miocene apes.

Soil erosion under forest hampers beech growth: Impacts of understory vegetation degradation by sika deer

The overpopulation of large herbivorous mammals degrades understory vegetation and hence promotes soil erosion in some northern hemisphere forests. Topsoil removal via erosion reduces soil water and nitrogen (N) content as well as exposes tree roots. However, whether topsoil removal inhibits tree growth through water and N uptake limitations has not been examined. This study aimed to measure the vertical length from exposed roots surface to the soil surface (exposed root height; ERH) under degraded forest understory vegetation and to evaluate its influence on tree growth. Specifically, we examined the relationships between ERH and beech growth and identified the timing of growth reduction based on annual tree-ring analysis at the individual scale. Furthermore, we discussed the effects of water and N limitations on beech growth reduction. We studied 12 Japanese beech trees on Mt. Sanpo, southern Kyushu Island, Japan, where understory vegetation was degraded and lost due to herbivory by Japanese sika deer from 1980 to 2003. Lower amounts of leaf production and relative growth rates of stem basal area increment (BAI) were observed in beech trees with higher ERH. Segmented regression analysis, which detects the changing time of a temporal trend (i.e., break-point), revealed that the BAI has resulted in a decreasing trend after 1997.6 ± 9.0 (yr), which coincided with the field observations on understory degradation in the study site. After the break-point of the BAI, the BAI of each study tree was negatively related to intrinsic water use efficiency, an integrated index of water limitation. These results suggest that soil erosion after the degradation of understory vegetation reduced beech growth, possibly because of the water limitations associated with root exposure. Reduced beech growth will reduce the supply of the litter layer that prevents topsoil erosion, which may enhance further soil erosion and tree growth reduction.

Herbivore–shrub interactions influence ecosystem respiration and biogenic volatile organic compound composition in the subarctic

Abstract. Arctic ecosystems are warming nearly 4 times faster than the global average, which is resulting in plant community shifts and subsequent changes in biogeochemical processes such as gaseous fluxes. Additionally, herbivores shape plant communities and thereby may alter the magnitude and composition of ecosystem respiration and biogenic volatile organic compound (BVOC) emissions. Here we determine the effect of large mammalian herbivores on ecosystem respiration and BVOC emissions in two southern and two northern sites in Swedish Scandes, encompassing mountain birch (LOMB) and shrub heath (LORI) communities in the south and low-herb meadow (RIGA) and shrub heath (RIRI) communities in the north. Herbivory significantly altered BVOC composition between sites and decreased ecosystem respiration at RIGA. The difference in graminoid cover was found to have a large effect on ecosystem respiration between sites as RIGA, with the highest cover, had 35 % higher emissions than the next highest-emitting site (LOMB). Additionally, LOMB had the highest emissions of terpenes, with the northern sites having significantly lower emissions. Differences between sites were primarily due to differences in exclosure effects and soil temperature and the prevalence of different shrub growth forms. Our results suggest that herbivory has a significant effect on trace gas fluxes in a productive meadow community and that differences between communities may be driven by differences in shrub composition.