Michael S. Bank

Loss of foundation species: consequences for the structure and dynamics of forested ecosystems

In many forested ecosystems, the architecture and functional ecology of certain tree species define forest structure and their species-specific traits control ecosystem dynamics. Such foundation tree species are declining throughout the world due to introductions and outbreaks of pests and pathogens, selective removal of individual taxa, and over-harvesting. Through a series of case studies, we show that the loss of foundation tree species changes the local environment on which a variety of other species depend; how this disrupts fundamental ecosystem processes, including rates of decomposition, nutrient fluxes, carbon sequestration, and energy flow; and dramatically alters the dynamics of associated aquatic ecosystems. Forests in which dynamics are controlled by one or a few foundation species appear to be dominated by a small number of strong interactions and may be highly susceptible to alternating between stable states following even small perturbations. The ongoing decline of many foundation species provides a set of important, albeit unfortunate, opportunities to develop the research tools, models, and metrics needed to identify foundation species, anticipate the cascade of immediate, short- and long-term changes in ecosystem structure and function that will follow from their loss, and provide options for remedial conservation and management.

Stable Isotope (N, C, Hg) Study of Methylmercury Sources and Trophic Transfer in the Northern Gulf of Mexico

We combined N, C, and Hg stable isotope measurements to identify the most important factors that influence MeHg accumulation in fish from the northern Gulf of Mexico (nGOM), and to determine if coastal species residing in the Mississippi River (MR) plume and migratory oceanic species derive their MeHg from the same, or different, sources. In six coastal species and two oceanic species (blackfin and yellowfin tuna), trophic position as measured by delta(15)N explained most of the variance in log[MeHg] (r(2) approximately 0.8), but coastal species and tuna fell along distinct, nearly parallel lines with significantly different intercepts. The tuna also had significantly higher delta(202)Hg (0.2-0.5 per thousand) and Delta(201)Hg ( approximately 1.5 per thousand) than the coastal fish (delta(202)Hg = 0 to -1.0 per thousand; Delta(201)Hg approximately 0.4 per thousand). The observations can be best explained by largely disconnected food webs rooted in different baseline delta(15)N signatures (MR-plume vs oceanic) and isotopically distinct MeHg sources, with oceanic MeHg having undergone substantial photodegradation ( approximately 50%) before entering the base of the food web. Given the MRs large, productive footprint in the nGOM and the potential for exporting prey and MeHg to the adjacent oligotrophic GOM, the disconnected food webs and different MeHg sources are consistent with recent evidence in other systems of important oceanic MeHg sources.

MERCURY BIOACCUMULATION AND TROPHIC TRANSFER IN SYMPATRIC SNAPPER SPECIES FROM THE GULF OF MEXICO

Consumption of marine fish is a major route of toxic methyl mercury (MeHg) exposure to ocean apex predators and human populations. Here we explore the influence of trophic structure on total mercury (Hg) accumulation in red snapper (RS, Lutjanus campechanus) and gray snapper (GS, Lutjanus griseus) from the coastal Louisiana region of the Gulf of Mexico, west of the Mississippi River. The objectives of this investigation were to: (1) determine the effectiveness of the use of offshore recreational fishing charter boats and marinas as sources of fish samples and (2) compare species differences in Hg bioaccumulation, trophic position, and carbon sources. Our data show that length-normalized Hg concentrations (> or = 97% as MeHg in tissue of both species) were 230% greater in GS in comparison to RS collected from the same general area. Stable C and N isotope signatures (delta15N and delta13C) indicate that GS occupy a slightly higher trophic position (approximately 30% of one trophic position higher) on the Gulf food web in comparison to RS and that GS appear to incorporate higher trophic positioned prey, continually and at smaller sizes. Mercury was strongly correlated with combined delta15N and delta13C in pooled species data, arguing that most of the substantial difference in Hg bioaccumulation between RS and GS can be explained by modest differences in their trophic position and, to a lesser degree, carbon sources, which had low variation and high overlap among species. These observations demonstrate that even minor to moderate differences in trophic position and food habits in sympatric species can create relatively large differences in bioaccumulation regimes and underscores the importance of quantitative characterization of trophic structure in marine MeHg bioaccumulation studies.

Temporal increase in organic mercury in an endangered pelagic seabird assessed by century-old museum specimens.

Methylmercury cycling in the Pacific Ocean has garnered significant attention in recent years, especially with regard to rising mercury emissions from Asia. Uncertainty exists concerning whether increases in anthropogenic emissions over time may have caused increased mercury bioaccumulation in the biota. To address this, we measured total mercury and, for a subset of samples, methylmercury (the bioaccumulated form of mercury) in museum feathers from an endangered seabird, the black-footed albatross (Phoebastria nigripes), spanning a 120-y period. We analyzed stable isotopes of nitrogen (δ15N) and carbon (δ13C) to control for temporal changes in trophic structure and diet. In post-1940 and -1990 feathers, we detected significantly higher mean methylmercury concentrations and higher proportions of samples exhibiting above deleterious threshold levels (∼40,000 ng·g−1) of methylmercury relative to prior time points, suggesting that mercury toxicity may undermine reproductive effort in the species. We also found higher levels of (presumably curator-mediated) inorganic mercury in older specimens of albatross as well as two nonpelagic species lacking historical exposure to bioavailable mercury, patterns suggesting that studies on bioaccumulation should measure methylmercury rather than total mercury when using museum collections. δ15N contributed substantially to models explaining the observed methylmercury variation. After simultaneously controlling for significant trends in δ13C over time and δ15N with methylmercury exposure, year remained a significant independent covariate with feather methylmercury levels among the albatrosses. These data show that remote seabird colonies in the Pacific basin exhibit temporal changes in methylmercury levels consistent with historical global and recent regional increases in anthropogenic emissions.

Spatial distribution of guanaco mating sites in southern Chile: conservation implications

We investigated the distribution, habitat characteristics and conservation value of mating areas in a wild population of guanacos in Torres del Paine National Park, Chile. Copulating guanacos were observed directly during December–January 1994–1995, 1995–1996 and 1997–1998. The timing of the breeding season was homogeneous across years. Copulation sites occurred in vega (ephemeral wetland) habitats (48.6%) greater than expected based on their relative availability (12.6%). A multivariate habitat analysis revealed that habitat composition of breeding habitats differed significantly from non-breeding areas (P < 0.001). Availability of trees and shrubs (breeding=5.5%; non-breeding=35.6%; P < 0.001) and grassland (breeding=34.6%; non-breeding =9.5%; P < 0.001) contributed most to the variation in habitat types among areas. Relative occurrence of pumas was significantly lower in breeding areas (P=0.04). The spatial arrangement of mating sites was similar among years suggesting that adult female and male guanacos exhibited site fidelity to breeding sites. Since vegas were important habitats for mating it is critical that natural resource managers consider the importance of these ephemeral wetland resources to ensure for proper reproduction and viability of local guanaco populations. # 2002 Elsevier Science Ltd. All rights reserved.

Meeting report: Methylmercury in marine ecosystems--from sources to seafood consumers.

Mercury and other contaminants in coastal and open-ocean ecosystems are an issue of great concern globally and in the United States, where consumption of marine fish and shellfish is a major route of human exposure to methylmercury (MeHg). A recent National Institute of Environmental Health Sciences–Superfund Basic Research Program workshop titled “Fate and Bioavailability of Mercury in Aquatic Ecosystems and Effects on Human Exposure,” convened by the Dartmouth Toxic Metals Research Program on 15–16 November 2006 in Durham, New Hampshire, brought together human health experts, marine scientists, and ecotoxicologists to encourage cross-disciplinary discussion between ecosystem and human health scientists and to articulate research and monitoring priorities to better understand how marine food webs have become contaminated with MeHg. Although human health effects of Hg contamination were a major theme, the workshop also explored effects on marine biota. The workgroup focused on three major topics: a) the biogeochemical cycling of Hg in marine ecosystems, b) the trophic transfer and bioaccumulation of MeHg in marine food webs, and c) human exposure to Hg from marine fish and shellfish consumption. The group concluded that current understanding of Hg in marine ecosystems across a range of habitats, chemical conditions, and ocean basins is severely data limited. An integrated research and monitoring program is needed to link the processes and mechanisms of MeHg production, bioaccumulation, and transfer with MeHg exposure in humans.

Mercury in litterfall and upper soil horizons in forested ecosystems in Vermont, USA

Mercury (Hg) is an atmospheric pollutant that, in forest ecosystems, accumulates in foliage and upper soil horizons. The authors measured soil and litterfall Hg at 15 forest sites (northern hardwood to mixed hardwood/conifer) throughout Vermont, USA, to examine variation among tree species, forest type, and soils. Differences were found among the 12 tree species sampled from at least two sites, with Acer pensylvanicum having significantly greater litterfall total Hg concentration. Senescent leaves had greater Hg concentrations if they originated lower in the canopy or had higher surface:weight ratios. Annual litterfall Hg flux had a wide range, 12.6 to 28.5 µg/m(2) (mean, 17.9 µg/m(2) ), not related to forest type. Soil and Hg pools in the Oi horizon (litter layer) were not related to the measured Hg deposition flux in litterfall or to total modeled Hg deposition. Despite having lower Hg concentrations, upper mineral soil (A horizons) had greater Hg pools than organic soil horizons (forest floor) due to greater bulk density. Significant differences were found in Hg concentration and Hg/C ratio among soil horizons but not among forest types. Overall, our findings highlight the importance of site history and the benefits of collecting litterfall and soils simultaneously. Observed differences in forest floor Hg pools were strongly correlated with carbon pools, which appeared to be a function of historic land-use patterns.

Forced dispersal of juvenile guanacos (Lama guanicoe): causes, variation, and fates of individuals dispersing at different times

We examined adult-juvenile conflict in the guanaco (Lama guanicoe). During spring, territorial males become increasingly aggressive toward all juveniles born the previous year and begin expelling them from family groups. In an apparent effort to reduce aggression, juveniles display submissive crouches when being observed, approached, or attacked by the territorial male. Therefore, we assessed the influence of juvenile submissive behavior on the timing of dispersal and also examined if dispersal time was related to survival and reproductive performance as adults. We also evaluated hypotheses regarding the evolution of juvenile mammalian dispersal in the context of if and how each may favor the forced dispersal of juvenile guanacos by territorial males. Juveniles generally dispersed in late spring and early summer, and a nearly equal proportion of females (n=46; 48%) and males (n=49; 52%) dispersed. More-submissive animals generally dispersed later than less-submissive animals. Juvenile sex and dispersal time were not related to survival. In contrast, juvenile sex and dispersal time were related to reproductive performance. The probability of reproducing was highest when juveniles dispersed early and decreased with increasing time in family groups prior to dispersal. The largest proportion of juveniles was forced to disperse during a 2-week interval following the peak of the breeding season. Competition for food resources is likely very intense at this juncture and territorial males may force older juveniles to disperse in order to divert food resources to younger neonates. Additionally, juveniles may be forced to disperse after territorial males mate their mothers to prevent lost mating opportunities, because females leave territories when their offspring disperse and possibly prior to mating with males. We conclude that the forced dispersal of juvenile guanacos by territorial males is ultimately driven by competition for food resources on territories. The timing of dispersal, however, may be tempered by the chronology of matings between territorial males and particular adult females, and/or genetic relatedness between territorial males and juveniles.

Predation of guanacos (Lama guanicoe) by southernmost mountain lions (Puma concolor) during a historically severe winter in Torres del Paine National Park, Chile

The effects of mountain lion Puma concolor predation on guanaco Lama guanicoe mortality was investigated during the historically severe winter of 1995 in Torres del Paine National Park, Chile. The 45 guanaco carcasses located represented 3% of the entire guanaco population of which 74% were mountain lion kills, 13% died from malnutrition, 2% died from fence entanglement and 11% died from unknown causes. Depleted bone marrow fat was observed in 20% of all carcasses. The number of guanaco deaths differed among sex and age classes, particularly in juveniles (< 1 year of age), which died more often than expected (P < 0.001). Fresh guanaco carcasses (n= 19) killed by mountain lions were located in tree and shrub habitats 79% of the time and were significantly greater than expected values based on the relative availability of these habitats (P < 0.001). Observations of the spatial distribution of mountain lion kills and guanaco mixed-sex groups were similar (P= 0.10), suggesting that mountain lions responded to winter migratory movements made by guanacos. Overall, adult guanacos experienced surprisingly lower levels of mortality (adult male = 1%, adult female = 2%) than expected, despite the severe winter conditions, while juveniles experienced higher levels of mortality (13%) than expected. In comparison to other years (1991–96), the severe winter of 1995 (i.e. this investigation) had the greatest total mortality for juveniles, although the proportion of deaths caused by mountain lion predation for juveniles was greatest during other winters.

Mercury bioaccumulation in green frog (Rana clamitans) and bullfrog (Rana catesbeiana) tadpoles from Acadia National Park, Maine, USA

Mercury contamination in the northeastern United States, including Acadia National Park (ANP; ME, USA), is well documented and continues to be a public health issue of concern. Mercury contamination of wild amphibians has received little attention, however, despite reports of worldwide population declines. Here, we report total Hg and methyl Hg (MeHg) concentrations for water, sediment, and green frog (Rana clamitans) and bullfrog (Rana catesbeiana) tadpoles (age, approximately one year) from ANP. Total Hg concentrations (mean+/-standard error) in green frog and bullfrog tadpoles were 25.1+/-1.5 and 19.1+/-0.8 ng/g wet weight, respectively. Mean total Hg was highest for green frog tadpoles sampled from the Schooner Head site (ANP, Bar Harbor, ME, USA), a small, semipermanent beaver pond where Ranavirus was detected during the summer of 2003 sampling period. Methyl Hg comprised 7.6 to 40% of the total Hg in tadpole tissue (wet-wt basis), and mean total Hg levels in tadpoles were significantly different among pond sites (n = 9). Total Hg in pond water was a significant predictor of tadpole total Hg levels. Dissolved organic carbon was a significant predictor of both total Hg and MeHg in water, and total Hg in water also was strongly correlated with MeHg in water. Of the nine pond ecosystems sampled at ANP 44% had a methylation efficiency (water MeHg to total Hg ratio) of greater than 10%, and 33% had total Hg levels in sediment that were approximately equal to or greater than the established threshold level effect concentration for freshwater sediments (0.174 mg/kg dry wt). Our data indicate that wetland food webs in ANP likely are susceptible to high levels of total Hg bioaccumulation and that methylation dynamics appear to be influenced by local abiotic and biotic factors, including disturbances by beavers and in situ water chemistry patterns. These findings may be important to National Park Service resource managers, especially considering the class I airshed status of ANP and the strong potential for negative effects to aquatic ecosystem structure and function from Hg pollution.