Small Mammalian Herbivores Research Articles

Size-biased compositional impacts of small mammal herbivores in an old-field plant community

Plant species height (mature height), which strongly correlates with other measures of size, is widely considered an important functional trait. Taller species contribute greatly to ecosystem function and are frequently identified as strong competitors. Additionally, important ecological processes like self-thinning scale with plant species size. In spite of the clear functional importance of species size, evidence for size-based plant community assembly rules has been scant. While herbivory can be species specific and there are reasons why herbivores may target larger plant species, it is unknown whether herbivory has size-biased impacts on species composition. In an old-field plant community, we compared plant species composition in herbivore exclusion (caged) and control plots over five years to test whether herbivore activity favors species at one end of the size spectrum. Across 5 years, smaller species were modestly, but consistently, underrepresented in caged plots compared to control plots. This underrepresentation of relatively smaller plant species in caged plots, where seedling herbivory was greatly diminished, was driven by lower small plant species richness and abundance in those plots; the abundance and richness of tall species did not differ across treatments. Our findings demonstrate that reducing herbivore activity in our study community modestly but consistently altered plant species composition, slightly favoring larger species. There was no evidence that larger species are preferentially targeted by herbivores. Some research suggests, although it will require further investigation, that seedling herbivory, regardless of the species eaten, produces vegetation gaps that small plant species are more successful in colonizing.

Tropical savanna small mammals respond to loss of cover following disturbance: A global review of field studies

Small-mammal faunas of tropical savannas consist of endemic assemblages of murid rodents, small marsupials, and insectivores on four continents. Small mammals in tropical savannas are understudied compared to other tropical habitats and other taxonomic groups (e.g., Afrotropical megafauna or Neotropical rainforest mammals). Their importance as prey, ecosystem engineers, disease reservoirs, and declining members of endemic biodiversity in tropical savannas compels us to understand the factors that regulate their abundance and diversity. We reviewed field studies published in the last 35 years that examined, mostly experimentally, the effects of varying three primary endogenous disturbances in tropical savanna ecosystems—fire, large mammalian herbivory (LMH), and drought—on abundance and diversity of non-volant small mammals. These disturbances are most likely to affect habitat structure (cover or concealment), food availability, or both, for ground-dwelling small mammalian herbivores, omnivores, and insectivores. Of 63 studies (included in 55 published papers) meeting these criteria from the Afrotropics, Neotropics, and northern Australia (none was found from southern Asia), 29 studies concluded that small mammals responded (mostly negatively) to a loss of cover (mostly from LMH and fire); four found evidence of increased predation on small mammals in lower-cover treatments (e.g., grazed or burned). Eighteen studies concluded a combination of food- and cover-limitation explained small-mammal responses to endogenous disturbances. Only two studies concluded small-mammal declines in response to habitat-altering disturbance were caused by food limitation and not related to cover reduction. Evidence to date indicates that abundance and richness of small savanna mammals, in general (with important exceptions), is enhanced by vegetative cover (especially tall grass, but sometimes shrub cover) as refugia for these prey species amid a “landscape of fear,” particularly for diurnal, non-cursorial, and non-fossorial species. These species have been called “decreasers” in response to cover reduction, whereas a minority of small-mammal species have been shown to be “increasers” or disturbance-tolerant. Complex relationships between endogenous disturbances and small-mammal food resources are important secondary factors, but only six studies manipulated or measured food resources simultaneous to habitat manipulations. While more such studies are needed, designing effective ones for cryptic consumer communities of omnivorous dietary opportunists is a significant challenge.

Effects of Inhabiting and Life Patterns on the UV Spectral Properties of Small Mammalian Herbivores’ Urine

It is well known that avian predators can use prey excretions such as urine and feces to track their prey, and the urine and feces of small mammalian herbivores can reflect ultraviolet (UV) light and emit fluorescent light. There are still some debates as to whether UV visibility of small mammalian herbivores’ urine is used as a hunting cue by avian raptors. Some studies in Europe have demonstrated that diurnal raptors are capable of utilizing these cues to target key prey species. However, researchers in Australia have argued that raptors do not use the UV visibility of urine while hunting. To our knowledge, there are no reports from Asia concerning the ultraviolet spectral characteristics of small mammal herbivores’ urine. This study examined the UV spectral properties of urine from 6 small mammal herbivores species by comparing the UV reflectance and fluorescence spectra of urine from small mammalian herbivores living in plateau meadows, plateau shrubs, open marshland, farmland, and semi-desert grassland in China. In addition, we compared the UV spectral properties of urine from ground-dwelling species of rodents and subterranean species to determine whether ultraviolet visibility of small mammal herbivores’ urine could be used as a visual signal by Asian vole-eating raptors. The results showed that: 1) the SC370 values of urine from four small mammal herbivores species were ordered as plateau pika (plateau meadow) > root voles (plateau bush) > reed voles (swampland) > Brandt’s vole (desert grassland); and 2) UV fluorescence peak intensity and the wavelengths of urine from ground-dwelling species (such as the root vole, plateau pika, or Brandt’s vole) were significantly higher than those of subterranean-dwelling species (mandarin vole and plateau zokor). These results indicate that UV visibility of small mammal herbivores’ urine may act as a visual cue for raptors.

Small mammal herbivores part 3: taking a dietary history and providing nutritional support

The unique dietary needs of exotic companion mammal herbivores has been thoroughly explored in this series of articles. The veterinary nurse can be well-equipped and is in an excellent position to take a detailed husbandry and nutritional history, which can help to identify nutritional disease and problems with dietary management that may contribute to future pathogenesis. Providing nutritional support in the hindgut fermenter inpatient or outpatient is centred around both restoring the negative energy balance (as the target species will rapidly enter a catabolic state) and providing enough fibre to stimulate gastrointestinal motility.

Embedding trait‐based ecology within indigenous knowledge to advance sustainable management of Tibetan rangeland

AbstractQuestionsTraditional pastoralists attribute high numbers of plateau pikas (Ochotona curzoniae) to impoverished soils that favor more and better forage for pika, suggesting a bottom‐up control of pika density. Conversely, government policies focus on excessive numbers of this small mammalian herbivore as the primary top‐down cause of degradation in Tibetan rangeland. Despite concerted campaigns to reduce pika abundance in recent decades, the sustainability of Tibetan rangelands remains uncertain.LocationAlpine meadows on the Tibetan Plateau.MethodsWe proposed a conceptual model based on indigenous knowledge that predicted pika numbers from soil condition and plant traits. At three alpine meadow sites lightly grazed by livestock, we tested whether spatial variation in pika burrow density could be explained by changes in the functional composition of the plant community attributable to species turnover and intraspecific trait variation associated with changes in soil fertility.ResultsDue primarily to intraspecific trait variation, changes in the functional composition of the meadow community accounted for 56%–68% of the spatial variation in pika density, changes in the proportion of plant functional groups for 62%–74%, and changes in edaphic conditions for 71%–82%. Greater pika density was associated with a decline in soil phosphorus availability and a lower‐growing vegetation profile enriched in both the quantity and quality of forage preferred by pikas.ConclusionsThese results, which are in accord with indigenous knowledge, suggest that compensating soil phosphorus losses and maintaining a well‐calibrated grazing rotation can better manage pika populations and improve the sustainability of Tibetan rangelands. By combining trait‐based ecology and traditional knowledge, our study provides new insight into both understanding the dynamic complexity of grazing regimes and managing rangeland sustainability.

Effect of the presence of plateau pikas on the ecosystem services of alpine meadows

Abstract. The activity of small mammalian herbivores influences grassland ecosystem services in arid and semi-arid regions. Plateau pika (Ochotona curzoniae) was considered to be a focal organism to investigate the effect of small mammalian herbivores on meadow ecosystem services in alpine regions. In this study, a home-range scale was used to measure the forage available to livestock, water conservation, carbon sequestration and soil nutrient maintenance (total nitrogen, phosphorus and potassium) in the topsoil layer, and a quadrat scale was used to assess the biodiversity conservation of alpine meadows. This study showed that the forage available to livestock and water conservation was 19 % and 16 % lower in the presence of plateau pikas than in their absence, and biodiversity conservation, carbon sequestration, soil nitrogen and phosphorus maintenance was 15 %, 29 %, 10 % and 8.9 % higher in the presence of plateau pikas than in their absence. In contrast, it had no impact on soil potassium maintenance of meadow ecosystems in alpine regions. The forage available to livestock, biodiversity conservation and soil nutrient maintenance of meadow ecosystems in alpine regions had maximum values as the disturbance intensity of plateau pikas increased; the water conservation tended to decrease linearly with the increasing disturbance intensity of plateau pikas. These results present a pattern of plateau pikas influencing the ecosystem services of meadow ecosystems in alpine regions, enriching our understanding of the small mammalian herbivores in relation to grassland ecosystem service.

Small mammal herbivores part 1: digestive system adaptations to a herbivorous diet

Nutritional disease is common in exotic companion mammals, and the unique dietary adaptations of herbivorous species only compounds this issue. The target species (including lagomorphs, and hystricomorph or caviomorph rodents) exhibit anatomical and physiological adaptations to a plant-based diet that is low in calories and protein, and high in fibre. Digestive tract adaptations such as hypselodont dentition to hindgut fermentation will be reviewed. Veterinary nurses are in an excellent position to counsel pet owners on the appropriate nutrition of their companion animals, and understanding these unique adaptations provides the necessary baseline knowledge to make recommendations.

Small but mighty: Impacts of rodent‐herbivore structures on carbon and nutrient cycling in arctic tundra

Abstract Understanding arctic ecosystem function is key to understanding future global carbon (C) and nutrient cycling processes. However, small mammal herbivores can have effects on ecosystems as structure builders and these effects have been underrepresented in the understanding of arctic systems. We examined the impact of small mammal structures (hay piles, runways, latrines) on soils and plants in three arctic tundra regions near Utqiaġvik, Toolik Lake, and Nome, Alaska. Our aims were to (1) examine how vole and lemming structures influence plant and soil nutrient pools and microbial processes, (2) determine if structure effects were similar across tundra system types, and (3) understand how changes in the abundance and cover of these structures during different phases of small mammal multi‐annual population cycles might influence biogeochemical cycling. In general, small mammal structures increased nitrogen (N) availability in soils, although effects varied by study region. Across study regions, hay piles were relatively uncommon (lowest % cover) but increased multiple soil N and P pools, C‐ and N‐acquiring enzyme activities, and leaf phosphorus (P) concentrations, with the specific nutrient variables and size of the effects varying by study region. Latrines had the second highest cover and influenced multiple C, N and P pools, but their effects were mainly observed within a single region. Lastly, runways had the highest % cover of all activity types but increased the fewest number of soil nutrient variables. We conclude that by influencing soil nutrient availability and biogeochemical cycling, small mammal structures can influence bottom‐up regulation of ecosystem function, particularly during the high phase of the small mammal population cycle. Future changes in these population cycles might alter the role of small mammals in the Arctic and have lasting effects on system processes. Read the free Plain Language Summary for this article on the Journal blog

Species provenance and traits mediate establishment and performance in an invaded grassland

Abstract In many invaded grasslands, dominant exotic species can produce large amounts of litter that modify local abiotic conditions and species' interactions. These novel conditions can reduce native species abundance and promote the persistence of exotic species, yet the strength of this disparity may be influenced by how consumer pressure interacts with litter accumulation. Consumers may exacerbate this disparity by preferentially targeting native species or by promoting heterogeneity in microhabitats due to their movement and small‐scale ground disturbances that favours fast growing exotic species. How species respond to litter accumulation and consumer pressure may depend on either evolutionary differences, whereby exotics species may benefit from a lack natural predators, or by functional differences, in which species' physiological traits may confer fitness advantages to low‐light conditions or herbivory or granivory pressure. We examined the impact of litter presence and small mammal herbivory on the establishment and reproduction of functionally diverse exotic versus native species seeded across sites that naturally vary in resource availability in an annual invaded California grassland. We assessed whether seed mass and leaf nitrogen content (LNC) were predictive of successful establishment and reproduction. Litter accumulation affected exotic and native species differently, with litter significantly decreasing native recruitment and reproduction, while exotics were largely unaffected. Small mammals had a slight positive effect on the establishment of native species when litter was present but did not influence exotic species. Regardless of species provenance, larger seeded species established at a higher density while species with lower leaf nitrogen content had a higher density of reproductive individuals. Native species that successfully established and reproduced were functionally different in LNC than the resident community, while successful exotic species were functionally more similar to the resident community in LNC. Our study demonstrates that exotic species outperformed native species regardless of the presence of litter or herbivory pressure. Without the removal or thinning of litter, it is likely that exotic species will continue to dominate, resulting in positive feedback that further favours the persistence of exotic species within this invaded grassland system. Read the free Plain Language Summary for this article on the Journal blog.

Utilisation of Woody Plants by the Cape Porcupine in Mesic Savannas in South Africa.

Herbivory plays a fundamental role in determining the structure of savannas. The impacts of small and medium-sized mammalian herbivores on trees in savannas remain poorly understood because most research attention focuses on large herbivores such as elephants whose destructive effects on trees can be pervasive at landscape scales. On the other hand, feeding activities of generalist herbivores such as Cape porcupines on woody plants can lead to tree mortality. This study investigated the utilisation of woody plants by the Cape porcupine in three mesic savanna sites in South Africa. We determined the woody plant diet of the porcupine for the early and late dry seasons at Roodeplaat Farm in Gauteng Province, and at Goss Game Farm and Bisley Valley Nature Reserve in KwaZulu-Natal Province. Thirty and twenty randomly located quadrats (30 m × 30 m) were laid at Roodeplaat and Goss, respectively, while 10 smaller quadrats (10 m × 10 m) were laid at Bisley. We measured stem diameter and the length and width of bark scars made by porcupines on stems of woody plants. We collected ten dung samples from each study site in the wet and dry seasons for quantification of woody material in porcupine diet. Porcupine foraging behaviour impacted different tree species at each site: Vachellia robusta at Roodeplaat, Spirostachys africana at Goss and Vachellia nilotica at Bisley. Each of these trees was dominant at each site. More scarring and tree mortality were recorded at Bisley with almost 70% tree sapling mortality occurring on trees that porcupine fed on. The size of bark scars was greater at Goss (P < 0.01) than at Roodeplaat and Bisley, which were similar. The area of bark damage on S. africana trees differed significantly by stem diameter size class (P = 0.007) and was greater for small stems (size class < 7.1 cm) than the larger stems (size classes 7.1-14 cm and 14.1-21). For all the study sites, dung samples revealed that woody material contributed over 80% of the porcupine diet during the dry season, but was lower at 35% during the wet season for Roodeplaat, although it was consistently high for Bisley at 79%. Porcupine foraging activities substantially contributed to tree mortality at each site. We posit that porcupine induced mortality on dominant tree species at each site may contribute to structural heterogeneity in woody plant vegetation in mesic savannas.

Large and small herbivores have strong effects on tundra vegetation in Scandinavia and Alaska.

Large and small mammalian herbivores are present in most vegetated areas in the Arctic and often have large impacts on plant community composition and ecosystem functioning. The relative importance of different herbivores and especially how their specific impact on the vegetation varies across the Arctic is however poorly understood.Here, we investigate how large and small herbivores influence vegetation density and plant community composition in four arctic vegetation types in Scandinavia and Alaska. We used a unique set of exclosures, excluding only large (reindeer and muskoxen) or all mammalian herbivores (also voles and lemmings) for at least 20 years.We found that mammalian herbivores in general decreased leaf area index, NDVI, and abundance of vascular plants in all four locations, even though the strength of the effect and which herbivore type caused these effects differed across locations. In three locations, herbivore presence caused contrasting plant communities, but not in the location with lowest productivity. Large herbivores had a negative effect on plant height, whereas small mammalian herbivores increased species diversity by decreasing dominance of the initially dominating plant species. Above‐ or belowground disturbances caused by herbivores were found to play an important role in shaping the vegetation in all locations.Synthesis: Based on these results, we conclude that both small and large mammalian herbivores influence vegetation in Scandinavia and Alaska in a similar way, some of which can mitigate effects of climate change. We also see important differences across locations, but these depend rather on local herbivore and plant community composition than large biogeographical differences among continents.

Spring phenology dominates over light availability in affecting seedling performance and plant attack during the growing season

Climate change can have important effects on plant performance by altering the relationship between spring temperature and other abiotic factors, such as light availability. Higher temperatures can advance plant phenology so that seedling germination takes place when days are shorter, and affect light availability for understory plants by altering the relative timing of seedling germination and canopy closure. To predict the effects of climate-induced changes in phenology and light availability on plant performance and species interactions during the growing season, we need to determine i) how effects of plant phenology on plant performance and the plant-associated community depend on light availability, and ii) to what extent effects of phenology and light availability on plant performance are direct vs. mediated by changes in the plant-associated community. We conducted a multifactorial field experiment to test for the effect of germination timing and light availability on Quercus robur seedling traits and performance, as well as attack by specialist plant pathogens, insects, and small mammals. Germination timing strongly affected seedling performance whereas light availability’s effects were limited. Likewise, germination timing strongly affected herbivore and pathogen attack, whereas light availability and its interaction with germination timing explained a minor part of the variation. Small mammals preferentially attacked later germinating seedlings, which strongly affected plant survival, while insect herbivores and pathogens did not mediate the effect of germination timing and light availability on plant performance. The results showed that the effect of germination timing can have greater influence than light availability on plant performance and plant attack, and that small mammal herbivores can play a larger role than diseases and insect herbivores in mediating the effect of spring phenology on plant performance. Together, these findings advance our understanding of the consequences of climate-induced changes in spring phenology and the abiotic environment on plant performance within a community context.

Species interactions across trophic levels mediate rainfall effects on dryland vegetation dynamics

AbstractArid ecosystems are strongly limited by water availability, and precipitation plays a major role in the dynamics of all species in arid regions, as well as the ecosystem processes that occur there. However, understanding how biotic interactions mediate long‐term responses of dryland ecosystems to rainfall remains very fragmented. We report on a unique large‐scale field experiment spanning 25 yr and three trophic levels (plants, small mammal herbivores, predators) in a dryland ecosystem in the northern Chilean Mediterranean Region where we assessed how biotic interactions influence the long‐term plant community responses to precipitation. As the most persistent ecological changes in dryland systems may result from changes in the structure, cover, and composition of the perennial vegetation, we emphasized the interplay between bottom‐up and top‐down controls of perennial plants in our analyses. Rainfall was the primary factor affecting the dynamics of, and interactions among, plants and small mammals. Ephemeral plant cover dynamics closely tracked short‐term annual rainfall, but seemed unaffected by top‐down controls (herbivory). In contrast, the response of the perennial plant cover to precipitation was mediated by (1) a complex interplay between subtle top‐down (herbivory) controls that become more apparent in the long‐term, (2) competition with ephemeral plants during wet years, and (3) an indirect effect of predators on subdominant shrubs and perennial herbs. This long‐term field experiment highlights how climate‐induced responses of arid perennial vegetation are influenced by interactions across trophic levels and temporal scales. In the face of global change, understanding how multi‐trophic controls mediate dryland vegetation responses to climate is essential to properly managing the conservation of biodiversity in arid systems.

Herbivore absence can shift dry heath tundra from carbon source to sink during peak growing season

In arctic tundra, large and small mammalian herbivores have substantial impacts on the vegetation community and consequently can affect the magnitude of carbon cycling. However, herbivores are often absent from modern carbon cycle models, partly because relatively few field studies focus on herbivore impacts on carbon cycling. Our objectives were to quantify the impact of 21 years of large herbivore and large and small herbivore exclusion on carbon cycling during peak growing season in a dry heath tundra community. When herbivores were excluded, we observed a significantly greater leaf area index as well as greater vascular plant abundance. While we did not observe significant differences in deciduous dwarf shrub abundance across treatments, evergreen dwarf shrub abundance was greater where large and small herbivores were excluded. Both foliose and fruticose lichen abundance were higher in the large herbivore, but not the small and large herbivore exclosures. Net ecosystem exchange (NEE) likewise indicated the highest carbon uptake in the exclosure treatments and lowest uptake in the control (CT), suggesting that herbivory decreased the capacity of dry heath tundra to take up carbon. Moreover, our calculated NEE for average light and temperature conditions for July 2017, when our measurements were taken, indicated that the tundra was a carbon source in CT, but was a carbon sink in both exclosure treatments, indicating removal of grazing pressure can change the carbon balance of dry heath tundra. Collectively, these findings suggest that herbivore absence can lead to changes in plant community structure of dry heath tundra that in turn can increase its capacity to take up carbon.

Exploring seedling-based aspen (Populus tremuloides) restoration near range limits in the Intermountain West, USA

With the effects of climate change expected to intensify over the coming century, land managers will require more proactive and novel approaches to conserve and restore threatened ecosystems. In the US Intermountain West, quaking aspen (Populus tremuloides Michx.) is a foundation species of considerable conservation interest. However, traditional silvicultural practices in the region focus exclusively on its vegetative regeneration potential, limiting restoration efforts to locations where aspen is already established, and precluding approaches such as afforestation and assisted migration. Planting nursery-grown aspen seedlings could overcome these limitations. This approach has become common for forest land reclamation and afforestation projects in the boreal forests of western North America, but has received little attention in the US Intermountain West to date. In this study we explored the potential for seedling-based aspen restoration in an Intermountain West landscape. We planted 7,200 nursery-grown aspen seedlings across 15 unirrigated plots at three high elevation (~2,800 m) sites in southwestern Utah, USA and monitored their growth and survival rates for two growing seasons. Overall seedling survival was only 10% by the end of the study, with 49% of mortality due to apparent drought stress, 38% due to small mammal herbivory, and 3% due to unknown causes. 79% of the surviving seedlings were located in just two plots, where higher levels of soil moisture were present during the driest summer months. A subsequent test of wood mulch to retain moisture in one of the plots increased seedling survival to 62%, compared to 0% in a non-mulched treatment. Together, these findings indicate that seedling-based aspen restoration in these habitats was limited primarily by site conditions. For this approach to become operational, additional study on seedling quality, site selection and site preparation will be necessary.

Long-term and interactive effects of different mammalian consumers on growth, survival, and recruitment of dominant tree species.

Throughout the world, numerous tree species are reported to be in decline, either due to increased mortality of established trees or reduced recruitment. The situation appears especially acute for oaks, which are dominant features of many landscapes in the northern hemisphere. Although numerous factors have been hypothesized to explain reductions in tree performance, vertebrate herbivores and granivores may serve as important drivers of these changes. Here, using data from 8‐ and 14‐year‐old exclosure experiments, we evaluated the individual and interactive effects of large and small mammalian herbivores on the performance of three widespread oak species in California—coast live oak (Quercus agrifolia), California black oak (Q. kelloggii), and Oregon white oak (Q. garryana). Although impacts varied somewhat by species and experiment, herbivory by black‐tailed deer (Odocoileus hemionus columbianus) reduced the height and survival of juvenile coast live oaks and altered their architecture, as well as reduced the abundance of black oak seedlings, the richness of woody species and the cover of nonoak woody species. Small mammals (Microtus californicus and Peromyscus maniculatus) had even more widespread effects, reducing the abundance of black oak seedlings and the height and cover of all three oak species. We also detected numerous interactions between small mammals and deer, with one herbivore having positive or negative effects on oak abundance and cover when the other herbivore was either present or absent. For example, deer often had negative effects on seedling abundance only when, or even more so when, small mammals were present. In summary, mammalian consumers play crucial roles in limiting oak recruitment by reducing seedling abundance, maintaining trees in stunted states, and preventing them from reaching sapling stages and becoming reproductive. Interactions between large and small mammals can also alter the intensity and direction of their effects on trees.

Small mammal herbivores mediate the effects of soil nitrogen and invertebrate herbivores on grassland diversity.

Simultaneous reductions in herbivore abundance and increases in nitrogen deposition have led to radical shifts in plant communities worldwide. While the individual impacts of these human‐caused disturbances are apparent, few studies manipulate both herbivory and N, nor differentiate among herbivore guilds, to understand contingencies in the ability of these drivers to affect producer diversity and productivity. As such, understanding how the main and combined effects of increasing soil N with declining herbivores may influence plant community structure and function is critical to better understand the future of grassland ecosystems under multiple global change drivers.In this study, we asked: (a) What are the main effects of small mammal herbivores, invertebrate herbivores, and soil N on plant community structure and function? and (b) Are the effects of invertebrate herbivores and soil N on plant community structure and function contingent on small mammal herbivory? We used a nested design, with invertebrate and soil N treatments nested within small mammal manipulations in an existing tallgrass prairie. We measured plant community structure by quantifying plant richness, evenness, diversity, and composition across two full growing seasons. We also recorded total aboveground biomass to quantify grassland productivity.We found that small mammal herbivores strongly shaped plant diversity, species composition, and productivity. Small mammal herbivores also mediated the effects of soil N and invertebrate herbivores on grassland community structure, but not composition or productivity. Small mammal reduction lowered plant species richness while increasing aboveground biomass and altering compositional similarity. Invertebrate herbivores, in the presence of small mammals, promoted plant dominance by reducing evenness without altering compositional similarity. Additionally, soil nitrogen addition reduced plant richness, but only when small mammals were reduced, and no effects were detected on compositional similarity or productivity.Our findings provide further evidence that temperate grasslands can be strongly influenced by consumers, and that consumers mediate the effects of resources as well as other consumer guilds on producer evenness and richness. Taken together, we provide evidence of strong contingencies in the drivers of grassland structure, with small mammals directly altering plant diversity as well as mediating the effects of soil nitrogen and invertebrate herbivory on plant richness and evenness. Therefore, we suggest it is imperative to consider how consumer guilds and resource types may interact to shape grassland plant communities.

Livestock grazing impacts on plateau pika (Ochotona curzoniae) vary by species identity

Livestock and small burrowing mammals are potential ecosystem engineers and major drivers of ecosystem processes. Small mammals, often considered competitors with livestock, have been reported to increase in recent years and have been identified for their negative impacts on ecosystem services such as plant production and soil carbon sequestration. However, these increases and impacts by small mammals have rarely been studied with respect to how they may be influenced by livestock grazing. Here we experimentally manipulated large enclosures on the alpine grassland of Qinghai-Tibetan plateau, each with different livestock assemblages: yak (Bos grunniens), Tibetan sheep (Ovis aries), mixed populations of yak and sheep, and a control plot containing no livestock to study the potential impacts of livestock on the dominant small mammalian herbivore in the region, the plateau pikas (Ochotona curzoniae). We found that during the period of peak vegetative growth all pika populations in livestock experimental plots increased compared with those in the control plots. Additionally, pika populations and active pika burrow density increased more in experimental plots containing yaks compared with the sheep-only experimental plots. These results provide evidence that livestock grazing can increase the risk of pika outbreaks, which has led to this species being classified as a pest and subject to widespread control measures. We suggest explicitly incorporating livestock species identity in addition to grazing intensity and duration into management considerations to ensure maintenance of native biodiversity and long-term sustainability on the alpine grasslands of the Qinghai-Tibetan plateau.

Neighbourhood crowding severely limits seed offspring recruitment in a temperate mesic old-field meadow

Testing the full impact of neighbourhood crowding within natural vegetation requires more than just effects incurred by established plants. It must also include measurements that take into account suppressive effects on the earliest plant life stages of resident individuals — seeds, their germination (emergence of radicles and cotyledons), and very young rooted seedlings. In this study, we explored the potential for these effects in a field experiment spanning three years, using a novel design for controlling granivory and small mammal herbivory. This allowed us to assess the limitations of natural crowding on seed recruitment success for non-resident species introduced into both natural and denuded neighbourhood plots within a temperate mesic old field meadow in eastern Ontario, Canada. Our results show that crowding by standing vegetation of resident species caused an overall reduction of seed recruitment success by more than 90%. These data provide strong inference that suppression resulting directly from near neighbour effects are likely to impose routinely intense natural selection within temperate mesic old field habitats like our study site. The consequences of this selection, in terms of traits promoting plant fitness under competition, are traditionally interpreted in terms of superior resource depletion/uptake, typically associated with greater growth accumulation and larger potential body size. We suggest, however, that these consequences are rare. Individuals of any species approach maximum potential body size only when near neighbour effects are relatively weak — not within crowded neighbourhoods. Recent studies suggest that severe neighbourhood crowding (where virtually all resident plants are forced to remain relatively small) selects instead for ‘reproductive economy’ — i.e., capacity to produce at least a few (or even at least one) offspring despite severe body size suppression, involving a relatively small minimum reproductive threshold size. Potential for additional component traits of reproductive economy are also suggested for investigation in future research.

Metagenomic sequencing provides insights into microbial detoxification in the guts of small mammalian herbivores (Neotoma spp.).

Microbial detoxification of plant toxins influences the use of plants as food sources by herbivores. Stephen's woodrats (Neotoma stephensi) specialize on juniper, which is defended by oxalate, phenolics and monoterpenes, while closely related N. albigula specialize on cactus, which only contains oxalate. Woodrats maintain two gut chambers harboring dense microbial communities: a foregut chamber proximal to the major site of toxin absorption, and a cecal chamber in their hindgut. We performed several experiments to investigate the location and nature of microbial detoxification in the woodrat gut. First, we measured toxin concentrations across gut chambers of N. stephensi. Compared to food material, oxalate concentrations were immediately lower in the foregut, while concentrations of terpenes remained high in the foregut, and were lowest in the cecal chamber. We conducted metagenomic sequencing of the foregut chambers of both woodrat species and cecal chambers of N. stephensi to compare microbial functions. We found that most genes associated with detoxification were more abundant in the cecal chambers of N. stephensi. However, some genes associated with degradation of oxalate and phenolic compounds were more abundant in the foregut chambers. Thus, microbial detoxification may take place in various chambers depending on the class of chemical compound.