Re-framing deer herbivory as a natural disturbance regime with ecological and socioeconomic outcomes in the eastern United States.

Natural disturbances are critical ecosystem processes, with both ecological and socioeconomic benefits and disadvantages. Large herbivores are natural disturbances that have removed plant biomass for millions of years, although herbivore influence likely has declined during the past thousands of years corresponding with extinctions and declines in distributions and abundances of most animal species. Nonetheless, the conventional view, particularly in eastern North America, is that herbivory by large wild herbivores is at unprecedented levels, resulting in unnatural damage to forests. Here, we propose consideration of large herbivores as a natural disturbance that also imparts many crucial ecological advantages, using white-tailed deer (Odocoileus virginianus), the only wild large herbivore remaining throughout the eastern U.S., as our focal species. We examined evidence of detrimental effects of browsing on trees and forbs. We then considered that deer contribute to both fuel reduction and ecological restoration of herbaceous plants and historical open forests of savannas and woodlands by controlling tree and shrub densities, mimicking the consumer role of fire. Similarly to other disturbances, deer disturbance 'regimes' are uneven in severity across different ecosystems and landscapes, resulting in heterogeneity and diversity. In addition to biodiversity support and fuel reduction, socioeconomic benefits include >$20 billion dollars per year by 10 million hunters that support jobs and wildlife agencies, non-consumptive enjoyment of nature by 80 million people, cultural importance, and deer as ecological ambassadors, whereas costs include about $5 billion and up to 450 human deaths per year for motor vehicle accidents, along with crop damage and disease transmission. From a perspective of historical ecology rather than current baselines, deer impart a fundamental disturbance process with many ecological benefits and a range of socioeconomic effects.

Multiple Browsers Structure Tree Recruitment in Logged Temperate Forests.

Historical extirpations have resulted in depauperate large herbivore assemblages in many northern forests. In eastern North America, most forests are inhabited by a single wild ungulate species, white-tailed deer (Odocoileus virginianus), and relationships between deer densities and impacts on forest regeneration are correspondingly well documented. Recent recolonizations by moose (Alces americanus) in northeastern regions complicate established deer density thresholds and predictions of browsing impacts on forest dynamics because size and foraging differences between the two animals suggest a lack of functional redundancy. We asked to what extent low densities of deer + moose would structure forest communities differently from that of low densities of deer in recently logged patch cuts of Massachusetts, USA. In each site, a randomized block with three treatment levels of large herbivores-no-ungulates (full exclosure), deer (partial exclosure), and deer + moose (control) was established. After 6-7 years, deer + moose reduced stem densities and basal area by 2-3-fold, Prunus pensylvanica and Quercus spp. recruitment by 3-6 fold, and species richness by 1.7 species (19%). In contrast, in the partial exclosures, deer had non-significant effects on stem density, basal area, and species composition, but significantly reduced species richness by 2.5 species on average (28%). Deer browsing in the partial exclosure was more selective than deer + moose browsing together, perhaps contributing to the decline in species richness in the former treatment and the lack of additional decline in the latter. Moose used the control plots at roughly the same frequency as deer (as determined by remote camera traps), suggesting that the much larger moose was the dominant browser species in terms of animal biomass in these cuts. A lack of functional redundancy with respect to foraging behavior between sympatric large herbivores may explain combined browsing effects that were both large and complex.

Ungulate browsers promote herbaceous layer diversity in logged temperate forests.

Ungulates are leading drivers of plant communities worldwide, with impacts linked to animal density, disturbance and vegetation structure, and site productivity. Many ecosystems have more than one ungulate species; however, few studies have specifically examined the combined effects of two or more species on plant communities. We examined the extent to which two ungulate browsers (moose [Alces americanus]) and white-tailed deer [Odocoileus virginianus]) have additive (compounding) or compensatory (opposing) effects on herbaceous layer composition and diversity, 5-6 years after timber harvest in Massachusetts, USA. We established three combinations of ungulates using two types of fenced exclosures - none (full exclosure), deer (partial exclosure), and deer + moose (control) in six replicated blocks. Species composition diverged among browser treatments, and changes were generally additive. Plant assemblages characteristic of closed canopy forests were less abundant and assemblages characteristic of open/disturbed habitats were more abundant in deer + moose plots compared with ungulate excluded areas. Browsing by deer + moose resulted in greater herbaceous species richness at the plot scale (169 m(2)) and greater woody species richness at the subplot scale (1 m(2)) than ungulate exclusion and deer alone. Browsing by deer + moose resulted in strong changes to the composition, structure, and diversity of forest herbaceous layers, relative to areas free of ungulates and areas browed by white-tailed deer alone. Our results provide evidence that moderate browsing in forest openings can promote both herbaceous and woody plant diversity. These results are consistent with the classic grazing-species richness curve, but have rarely been documented in forests.

A climatic driver for abrupt mid-Holocene vegetation dynamics and the hemlock decline in New England.

The mid-Holocene decline of eastern hemlock is widely viewed as the sole prehistorical example of an insect- or pathogen-mediated collapse of a North American tree species and has been extensively studied for insights into pest-host dynamics and the consequences to terrestrial and aquatic ecosystems of dominant-species removal. We report paleoecological evidence implicating climate as a major driver of this episode. Data drawn from sites across a gradient in hemlock abundance from dominant to absent demonstrate: a synchronous, dramatic decline in a contrasting taxon (oak); changes in lake sediments and aquatic taxa indicating low water levels; and one or more intervals of intense drought at regional to continental scales. These results, which accord well with emerging climate reconstructions, challenge the interpretation of a biotically driven hemlock decline and highlight the potential for climate change to generate major, abrupt dynamics in forest ecosystems.