Edward K. Faison

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.

Postglacial climate reconstruction based on compound‐specific D/H ratios of fatty acids from Blood Pond, New England

We determined hydrogen isotope ratios of individual fatty acids in a sediment core from Blood Pond, Massachusetts, USA, in order to reconstruct climate changes during the past 15 kyr. In addition to palmitic acid (C16n-acid), which has been shown to record lake water D/H ratios, our surface sediments and down core data indicate that behenic acid (C22n-acid), produced mainly by aquatic macrophytes, is also effective for capturing past environmental change. Calibration using surface sediments from two transects across eastern North America indicates that behenic acid records δD variation of lake water. Down core variations in δD values of behenic acid and pollen taxa are consistent with the known climate change history of New England. By evaluating the hypothesis that D/H fractionations of long chain even numbered fatty acids (C24-C32n-acids) relative to lake water provide independent estimates of relative humidity during the growing season, we find that differences between lake-level records and isotopically inferred humidity estimates may provide useful insight into seasonal aspects of the hydrologic cycle. Combined analyses of D/H of short and long chain fatty acids from lake sediment cores thus allow reconstructions of both past temperature and growing season relative humidity. Comparison of δD records from two lakes in New England provides critical information on regional climate variation and abrupt climate change, such as the 8.2 ka event.

LONG‐TERM HISTORY OF VEGETATION AND FIRE IN PITCH PINE–OAK FORESTS ON CAPE COD, MASSACHUSETTS

Human disturbance in northeastern North America over the past four cen- turies has led to dramatic change in vegetation composition and ecosystem processes, obscuring the influence of climate and edaphic factors on vegetation patterns. We use a paleoecological approach on Cape Cod, Massachusetts, to assess landscape-scale variation in pitch pine-oak vegetation and fire occurrence on the pre-European landscape and to determine changes resulting from European land use. Fossil pollen and charcoal preserved in seven lakes confirm a close link between landform and the pre-European distribution of vegetation. Pine forests, dominated by Pinus rigida, were closely associated with xeric outwash deposits, whereas oak-hardwood forests were associated with landforms having finer grained soils and variable topography. In general, fire was much more abundant on Cape Cod than most other areas in New England, but its occurrence varied geographically at two scales. On the western end of Cape Cod, fires were more prevalent in pine forests (outwash) than in oak-hardwood forests (moraines). In contrast, fires were less common on the narrow and north-south trending eastern Cape, perhaps because of physical limits on fire spread. The most rapid and substantial changes during the past 2000 years were initiated by European settlement, which produced a vegetation mosaic that today is less clearly tied to landform. Quercus and other hardwood trees declined in abundance in the early settlement period in association with land clearance, whereas Pinus has increased, especially during the past century, through natural reforestation and planting of abandoned fields and pastures. An increase in fossil charcoal following European settlement suggests that fire occurrence has risen substantially as a result of forest clearance and other land uses, reaching levels greater than at any time over the past 2000 years. Although fire was undoubtedly used by Native Americans and may have been locally important, we find no clear evidence that humans extensively modified fire regimes or vegetation before European settlement. Instead, climate change over the past several thousand years and European land use over the past 300 years have been the most important agents of change on this landscape.

Moose Foraging in the Temperate Forests of Southern New England

Abstract Moose have recently re-colonized the temperate forests of southern New England, raising questions about this herbivores effect on forest dynamics in the region. We quantified Moose foraging selectivity and intensity on tree species in relation to habitat features in central Massachusetts. Acer rubrum (Red Maple) and Tsnga canadensis (Eastern Hemlock) were disproportionately browsed; Pinus strobus (White Pine) was avoided. Foraging intensity correlated positively with elevation, distance to development, and watershed type and negatively with time since forest harvest, explaining 26% of the variation. Moose may interact with forest harvesting to contribute to a decline in Red Maple and Eastern Hemlock and an increase in White Pine in intensively browsed patches. Nonetheless, foraging impacts may diminish over time, as increasing temperatures and sprawling development increasingly restrict suitable Moose habitat.

Black Bear Foraging in Response to Beech Bark Disease in Northern Vermont

Abstract Several studies have demonstrated the relationship between black bear (Ursus americanus) reproductive success and the availability of hard mast. Beech bark disease (BBD) has been shown to reduce mast production in beech trees; however, no studies have investigated the relationship between bear foraging preferences of beech trees and the presence of BBD. To investigate this, we measured the severity of Nectria fungus and beech scale insect on freshly clawed and unclawed trees in pooled beech stands in northern Vermont. Our results suggest that bears do not have a strong preference for climbing healthier beech trees in aftermath forests. Changes in landscape composition and in cultural values appear to have been more important processes affecting black bear populations today than has BBD; however, we still advocate management strategies to minimize the impacts of this disease.

Ungulate browsers promote herbaceous layer diversity in logged temperate forests

Abstract 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 m2) and greater woody species richness at the subplot scale (1 m2) 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.

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.

To sprout or not to sprout: Multiple factors determine the vigor of Kalmia latifolia (Ericaceae) in southwestern Connecticut

Abstract Kalmia latifolia has declined in southern New England and other parts of its range in recent decades. This long-term decline is generally attributed to abiotic forces (i.e., low light levels in maturing forests) with little attention to the possible role that top-down effects from ungulate herbivory may be playing. We examined the extent to which mature K. latifolia is capable of sprouting under a relatively undisturbed forest canopy—both after severe stem injury and when uninjured—and tested the hypothesis that, in areas with high deer densities, herbivory may exceed abiotic forces in controlling the dynamics of K. latifolia. A block design experiment with white-tailed deer (Odocoileus virginianus) exclusion and control as treatments and landscape position (hilltop and low slope) as block was established in 2008. Canopy openness was measured in the two treatments within each block using hemispherical canopy photos. Survival and sprouting vigor of cut and uncut K. latifolia stems were monitored over 4 years and analyzed using Bayesian Information Criteria model selection with deer herbivory, percent canopy openness, and slope position as predictor variables. Canopy openness and slope position were important drivers of adult K. latifolia survival and sprouting capacity, whereas deer herbivory and slope position were the most important drivers of sprouting vigor on cut stems. Our results suggest that in a relatively undisturbed forest with high deer densities, herbivory does not exceed abiotic factors in determining adult K. latifolia vigor over the short term, but herbivory and slope position are more important than light in determining sprouting vigor after stem cutting.

Long-Term Deer Exclusion Has Complex Effects On A Suburban Forest Understory

Abstract Herbivory by deer is one of the leading biotic disturbances on forest understories (i.e., herbs, small shrubs, and small tree seedlings). A large body of research has reported declines in height, abundance, and reproductive capacity of forbs and woody plants coupled with increases in abundance of graminoids, ferns, and exotic species due to deer herbivory. Less clear is the extent to which (and the direction in which) deer alter herbaceous layer diversity, where much of the plant diversity in a forest occurs. We examined the effect of 15 y of deer exclusion on the understory of a suburban hardwood forest in Connecticut exposed to decades of intensive herbivory by white-tailed deer (Odocoileus virginianus). We compared species richness (at subplot and plot scale), individual species and life form group abundance (% cover), and community composition between grazed and exclosure plots, as well as between mesic and wet soil blocks. Forb cover was more than twice as abundant in exclosure as in grazed plots, whereas sedge (Carex spp.) cover was 28 times more abundant, and exotic species cover generally higher in grazed than in exclosure plots. Native and exotic species richness were both higher in grazed than exclosure plots at the subplot scale, and native herbaceous richness was higher in grazed plots at both spatial scales. In contrast, native shrub richness increased with deer exclusion at the plot scale. Our results suggest that deer exclusion had contrasting effects on species richness, depending on plant life form, but that overall richness of both exotic and native plants declined with deer exclusion. In addition, site heterogeneity remained an important driver of vegetation dynamics even in the midst of high deer densities.