Showing papers in "Ecological Monographs in 2004"

Quantitative fatty acid signature analysis: a new method of estimating predator diets

Abstract: Accurate estimates of the diets of predators are required in many areas of ecology, but for many species current methods are imprecise, limited to the last meal, and often biased. The diversity of fatty acids and their patterns in organisms, coupled with the narrow limitations on their biosynthesis, properties of digestion in monogastric animals, and the prevalence of large storage reservoirs of lipid in many predators, led us to propose the use of quantitative fatty acid signature analysis (QFASA) to study predator diets. We present a statistical model that provides quantitative estimates of the proportions of prey species in the diets of individual predators using fatty acid signatures. We conducted simulation studies using a database of 28 prey species (n = 954 individuals) from the Scotian Shelf off eastern Canada to investigate properties of the model and to evaluate the reliability with which prey could be distinguished in the model. We then conducted experiments on grey seals (Halichoerus grypus, n = 25) and harp seals (Phoca groenlandica, n = 5) to assess quantitative characteristics of fatty acid deposition and to develop calibration coefficients for individual fatty acids to account for predator lipid metabolism. We then tested the model and calibration coefficients by estimating the diets of experimentally fed captive grey seals (n = 6, switched from herring to a mackerel/capelin diet) and mink kits (Mustela vison, n = 46, switched from milk to one of three oil-supplemented diets). The diets of all experimentally fed animals were generally well estimated using QFASA and were consistent with qualitative and quantitative expectations, provided that appropriate calibration coefficients were used. In a final case, we compared video data of foraging by individual free- ranging harbor seals (Phoca vitulina, n = 23) fitted with Crittercams and QFASA estimates of the diet of those same seals using a complex ecosystem-wide prey database. Among the 28 prey species in the database, QFASA estimated sandlance to be the dominant prey species in the diet of all seals (averaging 62% of diet), followed primarily by flounders, but also capelin and minor amounts of other species, although there was also considerable individual variability among seals. These estimates were consistent with video data showing sandlance to be the predominant prey, followed by flatfish. We conclude that QFASA provides estimates of diets for individuals at time scales that are relevant to the ecological processes affecting survival, and can be used to study diet variability within individuals over time, which will provide important opportunities rarely possible with other indirect methods. We propose that the QFASA model we have set forth will be applicable to a wide range of predators and ecosystems.

Rapid evolution of an invasive plant

Abstract: Exotic plants often face different conditions from those experienced where they are native. The general issue of how exotics respond to unfamiliar environments within their new range is not well understood. Phenotypic plasticity has historically been seen as the primary mechanism enabling exotics to colonize large, environmentally diverse areas. However, new work indicates that exotics can evolve quickly, suggesting that contemporary evolution may be more important in invasion ecology than previously appreciated. To determine the influence of contemporary evolution, phenotypic plasticity, and founder ef- fects in affecting phenotypic variation among introduced plants, we compared the size, fecundity, and leaf area of St. John's wort ( Hypericum perforatum) collected from native European and introduced western and central North American populations in common gardens in Washington, California, Spain, and Sweden. We also determined genetic rela- tionships among these plants by examining variation in amplified fragment length poly- morphism (AFLP) markers. There was substantial genetic variation among introduced populations and evidence for multiple introductions of H. perforatum into North America. Across common gardens in- troduced plants were neither universally larger nor more fecund than natives. However, within common gardens, both introduced and native populations exhibited significant lat- itudinally based clines in size and fecundity. Clines among introduced populations broadly converged with those among native populations. Introduced and native plants originating from northern latitudes generally outperformed those originating from southern latitudes when grown in northern latitude gardens of Washington and Sweden. Conversely, plants from southern latitudes performed best in southern gardens in Spain and California. Clinal patterns in leaf area, however, did not change between gardens; European and central North American plants from northern latitudes had larger leaves than plants from southern latitudes within these regions in both Washington and California, the two gardens where this trait was measured. Introduced plants did not always occur at similar latitudes as their most closely related native progenitor, indicating that pre-adaptation (i.e., climate matching) is unlikely to be the sole explanation for clinal patterns among introduced populations. Instead, results suggest that introduced plants are evolving adaptations to broad-scale environmental conditions in their introduced range.

Late‐quaternary vegetation dynamics in north america: scaling from taxa to biomes

Abstract: This paper integrates recent efforts to map the distribution of biomes for the late Quaternary with the detailed evidence that plant species have responded individual- istically to climate change at millennial timescales. Using a fossil-pollen data set of over 700 sites, we review late-Quaternary vegetation history in northern and eastern North America across levels of ecological organization from individual taxa to biomes, and apply the insights gained from this review to critically examine the biome maps generated from the pollen data. Higher-order features of the vegetation (e.g., plant associations, physiog- nomy) emerge from individualistic responses of plant taxa to climate change, and different representations of vegetation history reveal different aspects of vegetation dynamics. Veg- etation distribution and composition were relatively stable during full-glacial times (21 000- 17 000 yr BP) (calendar years) and during the mid- to late Holocene (7000-500 yr BP), but changed rapidly during the late-glacial period and early Holocene (16 000-8 000 yr BP) and after 500 yr BP. Shifts in plant taxon distributions were characterized by individ- ualistic changes in population abundances and ranges and included large east-west shifts in distribution in addition to the northward redistribution of most taxa. Modern associations such as Fagus-Tsuga and Picea-Alnus-Betula date to the early Holocene, whereas other associations common to the late-glacial period (e.g., Picea-Cyperaceae-Fraxinus-Ostrya/ Carpinus) no longer exist. Biomes are dynamic entities that have changed in distribution, composition, and structure over time. The late-Pleistocene suite of biomes is distinct from those that grew during the Holocene. The pollen-based biome reconstructions are able to capture the major features of late-Quaternary vegetation but downplay the magnitude and variety of vegetational responses to climate change by (1) limiting apparent land-cover change to ecotones, (2) masking internal variations in biome composition, and (3) obscuring the range shifts and changes in abundance among individual taxa. The compositional and structural differences between full-glacial and recent biomes of the same type are similar to or greater than the spatial heterogeneity in the composition and structure of present-day biomes. This spatial and temporal heterogeneity allows biome maps to accommodate in- dividualistic behavior among species but masks climatically important variations in taxo- nomic composition as well as structural differences between modern biomes and their ancient counterparts.

Functional strategies of chaparral shrubs in relation to seasonal water deficit and disturbance

Abstract: The study of interspecific variation in plant ecological strategies has revealed suites of traits associated with leaf life span and with maximum levels of water deficit (measured as leaf water potentials). Here, the relationship between these sets of traits was examined in a study of 20 co-occurring chaparral shrubs that vary in leaf habit, rooting depth, and regeneration strategies. Leaf life span (LLS) and minimum seasonal water potentials (ψmin) were not significantly correlated, suggesting that they are associated with independent aspects of functional variation. Multiple regression analyses of a large suite of physiological, functional, and phenological attributes in relation to these two “anchor traits” supported this view. Short LLS was significantly associated with high specific leaf area, high carbon assimilation and leaf nitrogen (per mass), early onset of growth, and a multistemmed, short stature growth form. This suite of traits was also associated with opportunistic regeneration following physical disturbance. Area-based gas exchange was not tightly linked to LLS. Low ψmin (i.e., greater water deficit) was associated with high wood density, small vessel diameters, thin twigs, low leaf area : sapwood area ratios, and early onset of leaf abscission. Among the evergreen species, this suite of traits was most characteristic of post-fire seeders, which depend on high drought tolerance for post-fire regeneration of seedlings. Plant stature was the only trait associated with both the LLS axis and the ψmin axis of functional variation. A two-dimensional strategy space, approximately defined by LLS and ψmin, can be used to distinguish contrasting strategies of drought tolerance vs. avoidance, and alternative modes of regeneration following fire and other disturbance. This conceptual scheme illustrates the strength of a trait-based approach to defining plant strategies in relation to resource availability and disturbance.

Seed mass and mast seeding enhance dispersal by a neotropical scatter-hoarding rodent

Abstract: Many tree species that depend on scatter-hoarding animals for seed dispersal produce massive crops of large seeds at irregular intervals. Mast seeding and large seed size in these species have been explained as adaptations to increase animal dispersal and reduce predation. We studied how seed size and seed abundance simultaneously influenced seed dispersal and predation by scatter-hoarding rodents in the large-seeded rain forest tree Carapa procera (Meliaceae) in French Guiana. We individually tracked the fates of 3000 seeds, using remote video monitoring and thread-marking. Seed size was manipulated by broadly varying intraspecific seed mass, whereas effects of seed abundance were examined by tracking seeds in three seed-rich years and two seed-poor years. The hypotheses, that seed mass and seed abundance both enhance dispersal success and that seed abundance reinforces the effect of seed mass, were supported by the results. Most seeds were removed by the scatter-hoarding rodent red acouchy (Myoprocta acouchy) and subsequently were buried in scattered, single-seeded caches up to distances >100 m. Seeds that were not removed failed to establish seedlings. Seed removal was slower, pre-removal seed predation was greater, and seed dispersal was less far in seed-rich years than in seed-poor years, suggesting poorer dispersal under seed abundance. However, this was more than counterbalanced by a disproportionally greater survival of cached seeds in seed-rich years. The per capita probability of seed survival and seedling establishment was at least 4½ times greater under seed abundance. Large seeds were removed faster, were more likely to be scatter-hoarded, and were dispersed farther away than smaller ones, resulting in a higher probability of seedling establishment for larger seeds. Size discrimination was greater under seed abundance, albeit only during seed removal. Overall, large seeds shed in rich years had the highest probability of seedling establishment. Hence, both larger seed size and greater seed abundance stimulate rodents to act more as dispersers and less as predators of seeds. We conclude that scatter-hoarding rodents can select for both large seed crops and large seeds, which may reinforce mast seeding.

An experimental test of the causes of forest growth decline with stand age

Abstract: The decline in aboveground wood production after canopy closure in even-aged forest stands is a common pattern in forests, but clear evidence for the mechanism causing the decline is lacking. The problem is fundamental to forest biology, commercial forestry (the decline sets the rotation age), and to carbon storage in forests. We tested three hypotheses about mechanisms causing the decline in wood growth by quantifying the complete carbon budget of developing stands for over six years (a full rotation) in replicated plantations of Eucalyptus saligna near Pepeekeo, Hawaii. Our first hypothesis was that gross primary production (GPP) does not decline with stand age, and that the decline in wood growth results from a shifft in partitioning from wood production to respiration (as tree biomass accumulates), total belowground carbon allocation (as a result of declining soil nutrient supply), or some combination of these or other sinks. An alternative hypothesis was that GPP declines with stand age and that the decline in aboveground wood production is proportional to the decline in GPP. A decline in GPP could be driven be reduced canopy leaf area and photosynthetic capacity resulting from increasing nutrient limitation, increased abrasion between tree canopies, lower turgor pressure to drive foliar expansion, or hydraulic limitation of water flux as tree height increases. A final hypothesis was a combination of the first two: GPP declines, but the decline in wood production is disproportionately larger because partitioning shifts as well. We measured the entire annual carbon budget (aboveground production and respiration, total belowground carbon allocation [TBCA], and GPP) from 0.5 years after seedling planting through 6 1/2 years (when trees were ~25m tall). The replicated plots included two densities of trees (1111 trees/ha and 10 000 trees/ha) to vary the ratio of canopy leaf mass to wood mass in the individual trees, and three fertilization regimes (minimal, intensive, and minimal followed by intensive after three years) to assess the role of nutrition in shaping the decline in GPP and aboveground wood production. The forest closed its canopy in 1-2 years, with peak aboveground wood production, coinciding with canopy closure, of 1.2-1.8 kg C.m-2yr-1. Aboveground wood production declined from 1.4 kg C.m-2yr-1 at age 2 to 0.60 kg C.m-2yr-1 at age 6. Hypothesis 1 failed: GPP declined from 5.0 kg C.m-2yr-1 at age 2 to 3.2 kg C.m-2yr-1 at age 6. Aboveground woody respiration declined from 0.66 kg C.m-2yr-1 at age 2 to 0.22 kg C.m-2yr-1 at age 6 and TBCA declined from 1.9 kg C.m-2yr-1 at age 2 to 1.4 kg C.m-2yr-1 at age 6. Our data supported hypothesis 3: the decline in aboveground wood production (42% of peak) was proportionally greater than the decline in canopy photosynthesis (64% of peak). The fraction of GPP partitioned to belowground allocation and foliar respiration increased with stand age and contributed to the decline in aboveground wood production. The decline in GPP was not caused by nutrient limitation, a decline in leaf area or in photsynthetic capacity, or (from a related study on the same site) by hydraulic limitation. Nutrition did interact with the decline in GPP and aboveground wood production, because treatments with high nutritient availablity declined more slowly than did our control treatment, which was fertilized only during stand establishment.

Multiple trait associations in relation to habitat differentiation among 17 floridian oak species

Abstract: Differentiation of species distributions along environmental gradients and phenotypic specialization help explain the co-presence of 17 oak species that might oth- erwise be expected to competitively exclude one another. In an effort to understand the role of niche differentiation in the co-presence of these congeneric species in north-central Florida, we examined the community structure of oak-dominated forests in this region in relation to environmental variables and a suite of life history and physiological traits. Landscape distribution patterns of oaks and other woody species were determined from randomly established plots in three state parks. Soil moisture, nutrient availability, and fire regime were found to be critical factors influencing community structure, and the distri- bution of oak species was strongly correlated with these gradients. Detrended correspon- dence analysis of species' distributions supported the grouping of oak species into three major community types: (1) hammock, (2) sandhill, and (3) scrub. Principal components of multiple traits also supported differentiation of oak species into these three groups. There is clear evidence for ecological sorting among the oaks in these forests as species partition environmental gradients more than expected by chance, and most functional traits of species correspond to species distributions as predicted based on a priori understanding of trait function. Across phylogenetic lineages, species showed evolutionary convergence in function and habitat preference. In contrast, leaf-level traits were conserved within phylogenetic lineages and were not well correlated with local habitat factors, but rather with the broader geographic distributions and northern range limits of species. The cu- mulative results reported here provide strong evidence that oak species specialize for par- ticular niches via trade-offs in functional traits, and such niche partitioning contributes to the high diversity of oak species at the landscape level. At smaller spatial scales, phylo- genetic diversity among the oaks is likely to be important in promoting their coexistence through other mechanisms.

Patterns of herbivory and decomposition in aquatic and terrestrial ecosystems

Abstract: Describing the relative magnitude and controls of herbivory and decomposition is important in understanding the trophic transference, recycling, and storage of carbon and nutrients in diverse ecosystems. We examine the variability in herbivory and decomposition between and within a wide range of aquatic and terrestrial ecosystems. We also analyze how that variability is associated with differences in net primary production and producer nutritional quality. Net primary production and producer nutritional quality are uncorrelated between the two types of system or within either type. Producer nutritional quality is correlated to the percentage of primary production consumed by herbivores or percentage of detrital production decomposed annually, regardless of whether the comparison is made between the two types of systems or within either type of system. Thus, producer nutritional quality stands out as a consistent indicator of the importance of consumers as top-down controls of producer biomass and detritus...

Oak forest carbon and water simulations: model intercomparisons and evaluations against independent data

Abstract: Models represent our primary method for integration of small-scale, process- level phenomena into a comprehensive description of forest-stand or ecosystem function. They also represent a key method for testing hypotheses about the response of forest ecosystems to multiple changing environmental conditions. This paper describes the eval- uation of 13 stand-level models varying in their spatial, mechanistic, and temporal com- plexity for their ability to capture intra- and interannual components of the water and carbon cycle for an upland, oak-dominated forest of eastern Tennessee. Comparisons between model simulations and observations were conducted for hourly, daily, and annual time steps. Data for the comparisons were obtained from a wide range of methods including: eddy covariance, sapflow, chamber-based soil respiration, biometric estimates of stand-level net primary production and growth, and soil water content by time or frequency domain reflectometry. Response surfaces of carbon and water flux as a function of environmental drivers, and a variety of goodness-of-fit statistics (bias, absolute bias, and model efficiency) were used to judge model performance. A single model did not consistently perform the best at all time steps or for all variables considered. Intermodel comparisons showed good agreement for water cycle fluxes, but considerable disagreement among models for predicted carbon fluxes. The mean of all model outputs, however, was nearly always the best fit to the observations. Not surprisingly, models missing key forest components or processes, such as roots or modeled soil water content, were unable to provide accurate predictions of ecosystem responses to short-term drought phenomenon. Nevertheless, an inability to correctly capture short-term physiolog- ical processes under drought was not necessarily an indicator of poor annual water and carbon budget simulations. This is possible because droughts in the subject ecosystem were of short duration and therefore had a small cumulative impact. Models using hourly time steps and detailed mechanistic processes, and having a realistic spatial representation of the forest ecosystem provided the best predictions of observed data. Predictive ability of all models deteriorated under drought conditions, suggesting that further work is needed to evaluate and improve ecosystem model performance under unusual conditions, such as drought, that are a common focus of environmental change discussions.

A neighborhood analysis of tree growth and survival in a hurricane-driven tropical forest

Abstract: We present a likelihood-based regression method that was developed to an- alyze the effects of neighborhood competitive interactions and hurricane damage on tree growth and survival. The purpose of the method is to provide robust parameter estimates for a spatially explicit forest simulator and to gain insight into the processes that drive the patterns of species abundance in tropical forests. We test the method using census data from the 16-ha Luquillo Forest Dynamics Plot in Puerto Rico and describe effects of the spatial configuration, sizes, and species of neighboring trees on the growth and survival of 12 dominant tree species representing a variety of life history strategies. Variation in size- dependent growth and mortality suggests a complex relationship between size, growth, and survival under different regimes of light availability. Crowding effects on growth and survival appear to be idiosyncratic to each individual species, and with the exception of pioneers, there is little commonality among species that share similar life histories. We also explain the implications of differential susceptibility to hurricane damage on species' growth and survival and on their ability to respond to damage to neighboring trees. Tree species in the Luquillo Forest Dynamics Plot differ strikingly in both their susceptibility to hurricane disturbance and the nature of their recovery from wind disturbance, through response of both adult plants and juveniles to enhanced resource availability. At the stand level, intense competitive thinning of densely packed saplings that grew after hurricane damage accounted for the majority of post-hurricane mortality, particularly for shade- intolerant species. At the individual species level, effects of previous hurricane damage on growth and survival depended primarily on variation in the quantity and quality of hurricane damage sustained by target species and their interaction with life history characteristics of these individual species. Finally, we compare models that make different assumptions about the effects of com- peting species on tree growth and survival (e.g., equivalence of competitors vs. distinct species-specific effects). Size effects alone could not account for growth and survival for the majority of target species. Our results also demonstrate that competing species have distinct per capita effects on growth of dominant target species. In contrast, we found moderate support for a model that assumed functional equivalence of competitors on sur- vival. Key words: competitive coefficient; equivalent competitors; growth, size-dependent; hurricane disturbance; Luquillo Forest Dynamics Plot, Puerto Rico; mortality, size-dependent; neighborhood effects; neutral theory.

Landscape and regional impacts of hurricanes in puerto rico

Abstract: Puerto Rico is subject to frequent and severe impacts from hurricanes, whose long-term ecological role must be assessed on a scale of centuries. In this study we applied a method for reconstructing hurricane disturbance regimes developed in an earlier study of hurricanes in New England. Patterns of actual wind damage from historical records were analyzed for 85 hurricanes since European settlement in 1508. A simple meteorological model (HURRECON) was used to reconstruct the impacts of 43 hurricanes since 1851. Long-term effects of topography on a landscape scale in the Luquillo Experimental Forest (LEF) were simulated with a simple topographic exposure model (EXPOS). Average return intervals across Puerto Rico for F0 damage (loss of leaves and branches) and F1 damage (scattered blowdowns, small gaps) on the Fujita scale were 4 and 6 years, respectively. At higher damage levels, a gradient was created by the direction of the storm tracks and the weakening of hurricanes over the interior mountains. Average ...

The influence of species diversity and stress intensity on community resistance and resilience

Abstract: Recent theory, such as the insurance hypothesis, suggests that higher species diversity may dampen perturbation dynamics within a community. The dynamics of a rocky intertidal macroalgal community were evaluated using an experimentally induced heat stress applied at the end of a 15-mo manipulation of diversity. This pulse event produced a gradient of thermal stress within plots and, consequently, different degrees of perturbation. In gen- eral, the resistance of the community to the thermal stress was forecast by the pre-stress cover of dominant species, total algal cover, and standing biomass. Because higher diversity treatments, especially those containing the dominant algal group, fucoids, had higher overall abundance, highest diversity treatments were the most severely affected. The stress was also relatively nonselective, in that species were reduced in roughly equivalent proportions, suggesting an important distinction for predicting when diversity will not influence dis- turbance dynamics. The resilience of the community was strongly dependent on which species were initially present in the plots and the degree of disturbance. In highly disturbed areas, although the recovery trajectory was similar in early successional stages, differences emerged later; these differences appear to be attributable to the composition of the sur- rounding regeneration pool. For treatments not receiving the thermal stress, low-diversity plots without fucoids remained in states unlike the reference condition for most of the monitored resilience period. But plots in high-diversity treatments, even areas within plots that had experienced moderate disturbance, returned to states similar to the reference quick- ly. Thus, resilience (but not resistance) results are consistent with the insurance hypothesis. Overall, diversity's influence on community dynamics is complex and will depend on the characteristics of the stress as well as the characteristics of the species present in the

A long-term study of competition and diversity of corals

Abstract: Variations in interspecific competition, abundance, and alpha and beta diversities of corals were studied from 1962 to 2000 at different localities on the reef at Heron Island, Great Barrier Reef, Australia. Reductions in abundance and diversity were caused by direct damage by storms and elimination in competition. Recovery after such reductions was influenced by differences in the size of the species pools of recruits, and in contrasting competitive processes in different environments. In some places, the species pool of coral larval recruits is very low, so species richness (S) and diversity (D) never rise very high. At other sites, this species pool of recruits is larger, and S and D soon rise to high levels. After five different hurricanes destroyed corals at some sites during the 38- year period, recovery times of S and D ranged from 3 to 25 years. One reason for the variety of recovery times is that the physical environment was sometimes so drastically changed during the hurricane that a long period was required to return it to a habitat suitable for corals. Once S and D have peaked during recolonization, they may either remain at a high level, or decline. In shallow water, with no deleterious changes in environmental conditions, S and D may not decline over time, because superior competitors cannot overtop inferior competitors without exposing themselves to deleterious aerial exposure at low tide. At other times and places, S and D did decline over time. One cause of this was a gradual deterioration of the physical environment, as corals grew upward into the intertidal region and died of exposure. S and D also fell because the wave action in hurricanes either killed colonies in whole or part, or changed the drainage patterns over the reef crest, leaving corals high and dry at low tide. At deeper sites, declines in S and D were sometimes caused by heavy wave action, or by interspecific competition, as some corals overgrew or overtopped their neighbors and eliminated them.

Species interaction strength: testing model predictions along an upwelling gradient

Abstract: A recent model predicts that species interactions in benthic marine communities vary predictably with upwelling regimes. To test this model, we studied the Pisaster–Mytilus interaction at 14 rocky intertidal sites distributed among three oceanographic regions along a 1300-km stretch of the U.S. West Coast. Regions included an intermittent-upwelling region (northern), a persistent-upwelling region (central), and a region of weak and infrequent upwelling (southern). We quantified predation rates by the sea star Pisaster ochraceus on its main prey Mytilus californianus by transplanting mussels into the sea star's low-zone foraging range and comparing the rate of mussel loss in +Pisaster plots to those in −Pisaster plots. To evaluate the relation between predation rates and key ecological processes and conditions, we quantified phytoplankton concentration and rates of mussel recruitment, mussel growth, mussel abundance, and sea star abundance. Predictions of the model are expressed as responses of predator an...

Fecundity of trees and the colonization–competition hypothesis

Abstract: Colonization-competition trade-offs represent a stabilizing mechanism that is thought to maintain diversity of forest trees. If so, then early-successional species should benefit from high capacity to colonize new sites, and late-successional species should be good competitors. Tests of this hypothesis in forests have been precluded by an inability to estimate the many factors that contribute to seed production and dispersal, particularly the many types of stochasticity that contribute to fecundity data. We develop a hierarchical Bayes modeling structure, and we use it to estimate fecundity schedules from the two types of data that ecologists typically collect, including seed-trap counts and observations of tree status. The posterior density is obtained using Markov-chain Monte Carlo techniques. The flexiblestructure yields estimatesofsizeandcovariateeffectsonseedproduction,variability associated with population heterogeneity, and interannual stochasticity (variability and se- rial autocorrelation), sex ratio, and dispersal. It admits the errors in data associated with the ability to accurately recognize tree status and process misspecification. We estimate year-by-year seed-production rates for all individuals in each of nine sample stands from two regions and up to 11 years. A rich characterization of differences among species and relationships among individuals allows evaluation of a number of hypotheses related to masting, effective population sizes, and location and covariate effects. It demonstrates large bias in previous methods. We focus on implications for colonization-competition and a related hypothesis, the successional niche—trade-offs in the capacity to exploit high re- source availability in early successional environments vs. the capacity to survive low- resource conditions late in succession. Contrary to predictions of trade-off hypotheses, we find no relationship between suc- cessional status and fecundity, dispersal, or expected arrivals at distant sites. Resultssuggest a mechanism for maintenance of diversity that may be more general than colonization- competition and successional niches. High variability and strong individual effects (vari- ability within populations) generate massive stochasticity in recruitment that, when com- bined with ''storage,'' may provide a stabilizing mechanism. The storage effect stabilizes diversity when species differences ensure that responses to stochasticity are not highly correlated among species. Process variability and individual effects mean that many species have the advantage at different times and places even in the absence of ''deterministic'' trade-offs. Not only does colonization vary among species, but also individual behavior is highly stochastic and weakly correlated among members of the same population. Although these factors are the dominant sources of variability in data sets (substantially larger than the deterministic relationships typically examined), they have not been not included in the models that ecologists have used to evaluate mechanisms of species coexistence (e.g., even individual-based models lack random individual effects). Recognition of the mechanisms of coexistence requires not only heuristic models that capture the principal sources of stochasticity, but also data-modeling techniques that allow for their estimation.

Geographic structure of adélie penguin populations: overlap in colony-specific foraging areas

Abstract: In an investigation of the factors leading to geographic structuring among Adelie Penguin (Pygoscelis adeliae) populations, we studied the size and overlap of colony- specific foraging areas within an isolated cluster of colonies. The study area, in the south- western Ross Sea, included one large and three smaller colonies, ranging in size from 3900 to 135 000 nesting pairs, clustered on Ross and Beaufort Islands. We used triangulation of radio signals from transmitters attached to breeding penguins to determine foraging loca- tions and to define colony-specific foraging areas during the chick-provisioning period of four breeding seasons, 1997-2000. Colony populations (nesting pairs) were determined using aerial photography just after egg-laying; reproductive success was estimated by com- paring ground counts of chicks fledged to the number of breeding pairs apparent in aerial photos. Foraging-trip duration, meal size, and adult body mass were estimated using RFID (radio frequency identification) tags and an automated reader and weighbridge. Chick growth was assessed by weekly weighing. We related the following variables to colony size: foraging distance, area, and duration; reproductive success; chick meal size and growth rate; and seasonal variation in adult body mass. We found that penguins foraged closest to their respective colonies, particularly at the smaller colonies. However, as the season pro- gressed, foraging distance, duration, and area increased noticeably, especially at the largest colony. The foraging areas of the smaller colonies overlapped broadly, but very little foraging area overlap existed between the large colony and the smaller colonies, even though the foraging area of the large colony was well within range of the smaller colonies. Instead, the foraging areas of the smaller colonies shifted as that of the large colony grew. Colony size was not related to chick meal size, chick growth, or parental body mass. This differed from the year previous to the study, when foraging trips of the large colony were very long, parents lost mass, and chick meals were smaller. In light of existing data on prey abundance in neritic waters in Antarctica suggesting that krill are relatively evenly distributed and in high abundance in the Southern Ross Sea, we conclude that penguins depleted or changed the availability of their prey, that the degree of alteration was a function of colony size, and that the large colony affected the location (and perhaps ultimately the size) of foraging areas for the smaller colonies. It appears, therefore, that foraging dynamics play a role in the geographic structuring of colonies in this species.

Feedback between individual host selection behavior and population dynamics in an eruptive herbivore

Abstract: We examined the role of population density in host selection behavior of an eruptive insect herbivore, the spruce beetle Dendroctonus rufipennis. We conducted field and laboratory experiments on spruce beetles from 29 endemic and eruptive populations in Alaska and Utah, USA, and Yukon, Canada. Beetles from both population phases colonized trees that had been felled to remove host defenses. However, only beetles from eruptive populations colonized defended, healthy trees. A series of laboratory assays using host-based media amended with varying concentrations of phytochemicals identified several factors affecting population-dependent responses to hosts. First, beetles were repelled by high concentrations of the predominant spruce monoterpene, alpha-pinene, but intermediate concentrations elicited entry and gallery construction. Second, heritability assays suggested high genetic variance of host selection behavior within populations, and between-population differences persisted following rearing in a common environment. Third, beetles from eruptive populations were less likely to enter medium amended with phytochemicals and constructed shorter galleries, which disagreed with our prediction and seemingly contradicted the field observations. However, fourth, beetle avoidance of high concentrations of alpha-pinene decreased with the addition of other beetles, and this effect was more pronounced among beetles from eruptive populations than among those from endemic populations. This interaction broadened the host range of eruptive beetles. We propose that such density-dependent behaviors can maintain heterogeneity among population phases and contribute to positive feedback in herbivore population dynamics. A conceptual model suggests how heritable and environmentally induced variation in host selection behavior can affect bimodal equilibria and numerical thresholds in eruptive species.

Population viability with fire in eryngium cuneifolium: deciphering a decade of demographic data

Abstract: We analyzed and modeled the demography of Eryngium cuneifolium ,a n herbaceous species endemic to the fire-prone Florida scrub, using 10 annual censuses (1990- 1999) of 11 populations at Archbold Biological Station. Nearly every aspect of the de- mography of this plant is affected by time since fire. Year, time since fire, life history stage, and plant age affected survival, growth, and fecundity of E. cuneifolium, but time since fire and life history stage had the most consistent effects. Survival, flowering stem pro- duction, and early seedling survival were highest in recently burned sites. Long-term sur- vival, growth, and fecundity were highest for yearling cohorts recruiting recently after fire, with the largest contrast between plants recruiting two years postfire and those recruiting more than a decade postfire. Prior (historical) stage also affected individual plant fates. For example, plants with prior stasis or regression in stage subsequently died in greater numbers than plants with prior advancement in stage. Historical analyses did not suggest any cost associated with the initiation of flowering. We used a matrix selection approach to explicitly model Eryngium cuneifolium popu- lation viability in relation to fire. This simulation strategy included preserving observed data and variances within projection matrices formed for individual combinations of pop- ulation and year. We built 54 of these matrices, each with six stages (seed bank, yearlings, vegetative plants, and three reproductive stages). Each of these matrices also represented a specific time since fire. In building matrices, we minimized the use of pooled data while retaining specific matrices whenever possible. In this way, we preserved both the correlation structure within individual matrices (populations, years) and protected patterns among ma- trices across the time-since-fire gradient. To deal with less-detailed data on recruitment processes, we evaluated 13 fertility and seed bank scenarios that bracketed a range of outcomes. All scenarios were similar in showing the positive effects of fire on the demography of E. cuneifolium. The scenario with high seed bank survival (0.5) and low germination rates (0-0.005) was the best predictor of observed postfire years of peak aboveground population size (;8 yr) and aboveground population disappearance (30-34 yr), and also did a good job of reproducing observed population trajectories. Finite rates of increase (l) were .1 only during the first decade postfire but then declined beyond a decade postfire. Although prior (historical) stage affected most individual de- mographic parameters, it did not significantly influence finite rates of increase. Elasticities were highest for stasis and germination from the seed bank. Elasticities for survival in- creased with time since fire, while growth and fertility elasticities decreased. In historical models (those with information on stage from the second-to-last year), the elasticities for stasis were higher and the elasticities for growth lower, compared to models without this history. Bootstrapping suggested small standard errors for several types of model output. Most matrix elements were positively correlated, suggesting that favorable conditions affect many life history stages similarly, and that simulations using element selection would provide a less conservative risk assessment than the matrix selection technique used. We used a stochastic simulation program to simulate changing demography with time since fire, with various fire-return intervals, and for various initial population sizes. We obtained estimates of extinction risk and probability of population decline. Even populations as large as thousands of individuals will become extinct in the absence of fire. Fire-return intervals of 15 yr or less are necessary for E. cuneifolium persistence at individual sites. Fires at intervals longer than 20 yr create substantial extinction risks, and intervals longer than 12 yr produce declining populations. Cycles of widely divergent, alternating short and long fire-return intervals caused slightly higher chances of extinction compared to regular fire-return intervals. Although shrub regrowth is implicated in the decreased viability of E. cuneifolium populations under regimes of infrequent fire, aboveground fuel increases are often too slow

Leaf demography and phenology in Amazonian rain forest : a census of 40,000 leaves of 23 tree species

Abstract: The periodicity, synchrony, and causes of variability in the demography of tree leaves in ecosystems with relatively aseasonal climates, such as tropical rain forests, is still poorly understood. To address this issue, we surveyed the timing of birth and death of .40 000 leaves of 1445 individuals of 23 evergreen rain forest species in several late primary and early secondary successional plant communities at San Carlos de Rio Negro, Venezuela, in the northern Amazon basin. In all species, the mortality rate generally in- creased with leaf age. However, in many species, deceleration of death rates with extreme leaf age was noted. In general, for each species, the age structure of leaf populations and the frequency distribution of leaf life span were broad. Species differed substantially in their leaf demography. Measured in their native habitats, seven species common to disturbed open sites had shorter median life spans (0.7 yr) than five species common to open but infertile Bana primary communities (1.9 yr average) or six species common to two tall primary forest communities (Tierra Firme and Caatinga), when measured in high-light conditions in the canopy (2.0 yr average). Variation in light availability had consistent effects on leaf life span in all species. Species native to Tierra Firme forest had average leaf life spans of 3.2, 1.9, and 1.6 yr, respectively, in deeply shaded understory microsites, in small gaps, and in sunlit mature tree canopies. Species native to Caatinga forest had average leaf life spans of 4.2, 3.4, and 2.5 yr, respectively, in these same microsite types. Two species common in gaps and in disturbed sites had much longer leaf life span in shaded understory than in open, disturbed microsites. For all species, responses were similar when trees were planted in sites differing in light availability, as when trees naturally established across light gradients. The rate of leaf production, the risk of leaf mortality, and the leaf life span were not periodic or related consistently to seasonality of climate. Negligible relationships existed between the mild annual dry season and either leaf production or leaf mortality in all species. Thus, leaf phenology and demography were essentially aseasonal in this tropical forest environment.

Quantifying scale in ecology: lessons from awave-swept shore

Abstract: Understanding the role of scale is critical to ecologists' ability to make appropriate measurements, to ''scale up'' from local, short-term experiments to larger areas and longer times, to formulate models of community structure, and to address important conservation problems. Although these issues have received increased attention in recent years, empirical measurements of the scales of ecologically important variables are still rare. Here, we measure the spatial and temporal scales of variability of 15 ecologically relevant physical and biological processes in the wave-swept intertidal zone at Mussel Point, near Hopkins Marine Station in California. We analyze temporal variability in wave height, ocean temperature, upwelling intensity, solar irradiance, and body temperature for periods ranging from ten minutes to fifty years. In addition, we measure spatial variation in shoreline topography, wave force, wave-induced disturbance, body temperature, species diversity, recruitment, primary productivity, and the abundances of grazers, predators, and the competitive dominant occupier of space. Each of these spatial variables is measured along three horizontal transects in the upper intertidal zone: a short transect (44 m long with sampling locations spaced at ;0.5-m intervals), a medium transect (175 m long with sampling locations spaced at ;1.7-m intervals), and a long transect (334 m long with sampling locations spaced at ;3.4-m intervals). Six different methods are used to quantify the scale of each variable. Distinct scales are evident in all but one of our temporal variables, demonstrating that our methods for quantifying scale can work effectively with relatively simple, periodic phenomena. However, our spatial results reveal basic problems that arise when attempting to measure the scale of variability for more complex phenomena. For a given variable and length of transect, different methods of calculating scale seldom agree, and in some cases estimates differ by more than an order of magnitude. For a given variable and method of calculating spatial scale, measurements are sensitive to the length of a transect; the longer the transect, the larger the estimate of scale. We propose that the ''1/ f noise'' nature of the data can explain both the variability among methods for calculating scale and the length dependence of spatial scales of variation, and that the 1/ f noise character of the data may be driven by the fractal geometry of shoreline topography. We conclude that it may not be possible to define a meaningful spatial scale of variation in this system. As an alternative to the boiled-down concept of ''scale,'' we suggest that it is more appropriate to examine explicitly the pattern in which variability changes with the extent of measurement (e.g., the spectrum). Knowledge of this pattern can provide useful ecological scaling ''rules'' even when a well-defined scale (or hierarchy of scales) cannot be discerned.

Synthesis of a six-year study of calcareous grassland responses to in situ co2 enrichment

Abstract: We exposed species-rich temperate calcareous grassland to elevated CO2 (600 μL/L) for six growing seasons and studied carbon (C) and nutrient (nitrogen [N] and phosphorus [P]) cycling, water relations, and plant community structure and diversity. CO2 enrichment stimulated leaf- and ecosystem-level daytime CO2 uptake and increased plant community productivity; relative CO2 effects on aboveground biomass were predicted with r 0.95) by precipitation prior to biomass harvests. The underlying mechanisms were water savings due to reduced leaf conductance under elevated CO2, allowing for more growth in this temporarily water-limited ecosystem; this effect was more important in dry years. At the plant-species level, no effects of [CO2] were found except for the subdominant sedges Carex flacca and C. caryophyllea, which responded positively to elevated CO2, mainly due to increased soil moisture. Bryophytes also responded to CO2 enrichment for the same reason. At a more aggregate level, elevated CO2 in...

Indirect effects of an introduced ungulate on pollination and plant reproduction

Abstract: Herbivores can affect plants not only directly through browsing and tram- pling, but also indirectly through other species. For example, herbivores could affect the interaction between plants and their pollinators. Because plant population density may affect plant-pollinator interactions and plant reproductive success, we hypothesized that herbi- vores could affect pollination and plant reproduction indirectly by modifying plant popu- lation density. Unlike previous hypotheses, which concerned individual-level effects on vegetative and reproductive traits, our hypothesis focuses on population-level effects and involves a feedback mechanism. To test this hypothesis, we conducted field studies in the temperate forest of the southern Andes, where introduced ungulates are a major source of anthropogenic alteration. For 10 animal-pollinated understory plants, we compared popu- lation density, pollinator visitation, pollen deposition in stigmas, and reproduction in four pairs of grazed and ungrazed sites. We found evidence of indirect effects of ungulates on pollination and reproduction only for the herb Alstroemeria aurea(Alstroemeriaceae). The general lack of evidence for indirect effects on most of the species may result from resistance to cattle browsing and trampling, or low statistical power. For A. aurea, we present additional evidence from trampling and hand-pollination experiments, plus path analyses of the effect of density on pollination and reproduction showing that: (1) cattle decrease the absolute and relative population density of this species through trampling; (2) density, particularly relative density, affects pollen deposition on stigmas; and (3) conspecific pollen deposition affects reproduction. Thus, our results indicate that, by directly reducing the population density of A. aurea, cattle are indirectly affecting its reproduction.

Cannibalism in a size‐structured population: energy extraction and control

Abstract: Recent size-structured cannibalistic models point to the importance of the energy gain by cannibals and also show that this gain may result in the emergence of giant individuals. We use a combination of a 10-year field study of a perch (Perca fluviatilis) population and quantitative within-season modeling of individual and population-level dy- namics to investigate which mechanisms are most likely to drive the dynamics of the studied perch population. We focused on three main aspects to explain observed discrepancies between earlier model predictions and data: (1) introduction of more than one shared resource between cannibals and victims, (2) whether or not several victim age cohorts are necessary to allow giant growth, and (3) the intensity of inter-cohort competition between young-of-the-year (YOY) perch and 1-yr-old perch. At the start of the study period, the perch population was dominated by ''stunted'' perch individuals, and recruitment of perch to an age of 1-yr-old was negligible. Following a major death in adult perch, strong recruitments of perch to 1-yr-old were thereafter observed for a number of years. As 1-yr-olds these successful recruiters subsequently starved to death due to competition with the new YOY. The few surviving adult perch accelerated substantially in growth and became ''giants.'' At the end of the study period, the perch population moved back to the situation with stunted individuals. There was a high agreement between observed diets of cannibalistic perch and those predicted by the model for both the stunted and the giant phases. Analyses of growth rates showed that cannibalistic perch could become giants on a diet of YOY perch only, but that a supplement with the second shared resource (macroinvertebrates) was needed to reach the observed sizes. Modeling of growth and diet in the giant phase showed an exploitative competitive effect of YOY perch on 1-yr-old perch, but a restriction in habitat use of 1-yr-old perch had to be assumed to yield the observed growth rate and diet. The resource dynamics of zooplankton and mac- roinvertebrates were both accurately predicted by the model. Also, YOY perch mortality was accurately predicted and, furthermore, suggested that one of the trawling methods used may underestimate the number of YOY perch when they increase in size. We conclude that the presence of a second shared resource and the restricted habitat use and absence of cannibalistic consumption by 1-yr-old perch individuals are two im- portant mechanisms to explain the discrepancy between model predictions and data. Our results also point to the fact that that the dynamics observed may be explained by complex dynamics not involving the presence of a giant and dwarf cycle.

Determinants of reproductive costs in the long-lived Black-legged Kittiwake: A multiyear experiment

Abstract: We studied reproductive costs of Black-legged Kittiwakes (Rissa tridactyla) in Prince William Sound, Alaska (USA) by removing entire clutches from randomly selected nests over four successive years, and then contrasting survival and fecundity of adults from manipulated and unmanipulated nests in each subsequent year. To elucidate mechanisms that lead to the expression of reproductive costs, we simultaneously characterized several behavioral and physiological parameters among adults in the two treatment groups. We also examined naturally nonbreeding adults that previously bred to determine their survival and future nonbreeding probabilities. Food availability varied during the study, being generally poor at the onset, and improving in later years. Adult nest attendance and body condition (assessed late in the chick- rearing period) varied accordingly among years, and between adults raising chicks and adults that had their eggs removed. Adults from unmanipulated nests incurred significant survival costs in all years, although fecundity costs were strongly expressed in only one of four years. Neither survival nor fecundity costs were strongly influenced by body condition or food availability, and no difference in reproductive costs was detected between the sexes. Although unmanipulated breeders survived at lower rates than manipulated breeders due to costs of reproduction, their survival rates were elevated compared to those of natural nonbreeders, presumably due to differences in individual ability. These findings indicate that models of adult survival must consider not only an organism's reproductive state, but also the factors that lead to that state. Although body condition appeared to be weakly related to survival, it was insufficient to explain the full magnitude of survival costs observed. We suggest that other parameters that were found to differ between treatment groups (e.g., rates of energy turnover, baseline levels of stress, and patterns of allocating body reserves) may be important mechanistic determinants of reproductive costs in kittiwakes, and potentially other long-lived species. Future efforts should move beyond simple assessments of body condition and toward more integrated measures of physiological condition when attempting to identify factors that influence how long-lived species balance the costs and benefits of reproduction.

Nothofagus regeneration dynamics in south‐central chile: a test of a general model

Abstract: Studies in Southern Hemisphere temperate forests have yielded a general model of how the successional status of shade-intolerant Nothofagus is related to site conditions and disturbance. According to model predictions, regeneration of most Nothofagus species is dependent on coarse-scale disturbance at favorable sites where shade-tolerant rain forest species co-occur. In contrast, at suboptimal sites where tree species richness is low, regeneration is not dependent on coarse-scale disturbance. We used variation in forest composition and structure as well as disturbance patterns in the forests of Puyehue National Park, Chile, to test several predictions from this general model. The study area presents a complex mosaic of disturbance patterns, including the effects of the strong 1960 earthquake as well as fire and wind disturbance. We reconstructed stand development on sites affected by these disturbances, and compared these patterns with the structure of old-growth forests along an elevation gradient from s...

Foliar nutrition, site quality, and temperature influence foliar chemistry of tallowwood (eucalyptus microcorys)

Abstract: The distribution of marsupial folivores in Australian eucalypt forests is extremely patchy and, although distribution patterns are known to be partly determined by nutritional factors, the role of plant secondary metabolites (PSMs) has been little studied. We investigated variation in the foliar concentrations of five constituents: total phenolics (TP); the foliar component bound by polyethylene glycol (PEGBC); the formylphloroglucinol compound, sideroxylonal (a strong deterrent of mammalian herbivory); the terpene, cineole; and nitrogen (N). These constituents were analyzed using near infrared spectroscopy in 501 individual tallowwood (Eucalyptus microcorys) trees at 42 sites across a broad geographic range in eastern Australia. We tested three hypotheses: (1) trees with lower foliar N produce higher PSM concentrations; (2) PSM concentrations are greater at colder sites; and (3) PSM concentrations are higher at sites with low resource availability. Hypothesis 1 was true for all PSMs considered and was the result of variation that occurred within sites. This is consistent with the carbon/nutrient balance hypothesis and with the hypothesis that PSMs protect leaves from photodamage. Hypothesis 2 was upheld for all constituents. This was consistent with both the growth–differentiation balance hypothesis, and with the hypothesis that PSMs represent an evolutionary adaptation to greater risks of photodamage and frost damage in cold environments. Hypothesis 3 was rejected, with one PSM, sideroxylonal, being affected by resource availability in a manner opposite to that predicted. The optimal defense hypothesis offers an explanation for this result (as sideroxylonal concentrations increased along with the apparent nutritional quality of the foliage, possibly to provide defense against higher rates of herbivory). We found that trees associated with koala fecal pellets had lower concentrations of sideroxylonal and cineole. We discuss this finding and further implications for the herbivores of Eucalyptus of the patterns of plant defense that we detected.

A new technique for maximum-likelihood canonical gaussian ordination

Abstract: Canonical correspondence analysis (CCA) is probably the most popular or- dination method in community ecology. However, it is only a heuristic approximation to maximum-likelihood estimated canonical Gaussian ordination (CGO), which is the ''ideal'' method. When proposed in the mid-1980s, CCA held two advantages over CGO: it was computationally cheaper, and its algorithm was not complex. However, an exponential increase in computing speed over the last two decades has meant that computation cost is no longer such a compelling advantage. The computational complexity of CGO has always been its major difficulty, even though it is statistically more sound and simpler to understand than CCA. For these reasons, no general computational framework or software has appeared until now. This article proposes a new class of statistical regression models called quadratic re- duced-rank vector generalized linear models (QRR-VGLMs) for maximum-likelihood es- timated CGO. This is achieved by extending a recently developed class of statistical models called RR-VGLMs. The extension is named QRR-VGLMs because of the addition of a quadratic form to each linear predictor, with the consequence that bell-shaped responses can be modeled as functions of latent environmental variables or gradients. QRR-VGLMs have several major positive features; for example, their framework is unifying and broad, so that canonical Gaussian ordination can potentially be performed on a wide range of data types. The two most important special cases of CGO in ecology, multispecies presence/ absence and Poisson abundance data, are considered in this article. The methodology is illustrated with a real data set using a software implementation written by the author in the S statistical language. The code, called the VGAM package in R, is object-oriented and free, and it allows QRR-VGLMs to be fitted to moderate-sized data sets conforming rea- sonably closely to the Gaussian model.

Limiting resources, disturbance, and diversity in phytoplankton communities

Abstract: Phytoplankton diversity, limiting resources, and disturbance were studied in two reservoirs, Eagle Mountain Lake (EML) and Joe Pool Lake (JPL), in north Texas, USA, for three summer growing seasons and two winters Availabilities of phosphorus, nitrogen, and silicon were measured as chemical concentrations, and availability of light was measured as irradiance during the photoperiod, averaged over the depth of the surface mixed layer A resource was defined as limiting whenever its availability fell below a threshold, treated as a parameter to be fitted Depth of the surface mixed layer and wind speed were taken as indices of disturbance associated with episodic mixing Hydrological disturbance was gauged by rainfall, inflow, and the variation of lake level We found that diversity was strongly and significantly correlated with the number of limiting resources in one lake (JPL), but not the other (EML) The onset of nitrogen and phosphorus limitation during the summer growing season is associated with increased phytoplankton diversity in JPL Regression modeling detected decreasing relationships between diversity and disturbance indices in one lake (JPL), but concave-down relationships in the other (EML), though the peaks of these relationships did not lie within the usual range of disturbance Predictive regressions for diversity incorporating both limiting resources and disturbance were partitioned into unique and shared effects of these factors There was a large unique effect of the number of limiting resources in JPL, but otherwise the shared effects were large relative to unique effects

Dynamics of subarctic wetland forests over the past 1500 years

Abstract: Boreal forests at high latitudes are climate-sensitive ecosystems that respond directly to environmental forcing by changing their position according to latitude or by changing their abundance at local and regional scales. South of the arctic treeline, external forcing (warming, cooling, drought, fire) necessarily results in the changing abundance of the impacted forests; in particular, the deforestation of well-drained sites through fire is the most important factor. In this study, we examined the changing abundance of wetland forests located at the arctic treeline (northern Quebec, Canada) during the last 1500 years, a period of known contrasting climatic conditions. Black spruce (Picea mariana) trees submerged in small lakes and peatland ponds and soil-peat stratigraphy were used con- currently to reconstruct the millennial-long developmental sequence of wetland stands as- sociated with moisture changes and fire disturbance. Changing lake levels from AD 300 to the present were identified based on radiocarbon-dated submerged paleosols and tree- ring cross-dating of submerged trees distributed in three wetlands from the same watershed. Dead and living trees in a standing position below and above present water level of a small lake (LE Lake) showed direct evidence of past water levels from the 12th century to the present day. Submerged subfossil trees from another lake (LB Lake) and two peatland ponds (PB Peatland) also responded synchronously to changes in soil moisture during the last 1500 years. Regional-scale catastrophic flooding around AD 1150, inferred from paleosol and subfossil tree data, eliminated riparian peat and wetland trees growing at least since AD 300. Also, the coincidence of events such as the mass mortality of wetland spruce and post-fire deforestation of a small hill surrounding LE Lake during the late 1500s suggests the impact of local-scale flooding, probably attributable to greater snow transportation and accumulation on the lake surface after fire disturbance. Massive tree mortality climaxed at ca. 1750, when all wetland trees at LB Lake and PB Peatland died because of permafrost disturbance and soil upthrusting. Lower water levels from AD 300 to 1750 were associated with drier conditions, possibly caused by greater evaporation and/or reduced snow accu- mulation. Permafrost development in shallow waters occurred during the Little Ice Age, after 1600. It is concluded that the climate at the eastern Canadian treeline was warmer and drier from AD 300 to the onset of the Little Ice Age and promoted tree establishment. The highest water levels were recorded recently (19th and 20th centuries), causing lake and peatland expansion. Any future moisture changes at these subarctic latitudes will result in important spatial rearrangements of wetland ecosystems.

A paleoecological perspective of vegetation succession on raised bog microforms

Abstract: Peat cores were taken from eight primary and one secondary raised bog sites throughout the United Kingdom in order to determine the pathways of succession between the microform (hummock, ridge, lawn, hollow, and pool) plant communities of the mid to late Holocene. All of these sites have undergone drainage and/or cutting in the recent past, at least. The plant communities were reconstructed by using plant macrofossil analysis, and they are defined primarily by the dominant species of Sphagnum. While plant macrofossils may be used to infer hydrological conditions, an alternative proxy, using testate amoebae, was utilized. This group of organisms was used to avoid circular arguments when relating the vegetation changes to hydrological events. Consequently, a conceptual model of late Holocene succession on British raised mires is constructed that elucidates succession between the microform plant communities according to changes in the water table depth, its stability, quality, and physical disturbance of the vegetation cover. The general sequence of ombrogenous communities was elucidated to be, with increasing height above the water tables, S. cuspidatum and S. auriculatum > Sphagnum sect. Cymbifolia taxa > Sphagnum sect. Acutifolia taxa > a return of Sphagnum sect. Cymbifolia taxa > non-Sphagnum mosses. A major pathway toward degradation of the mire is apparent in modern times, and recovery is typically via a phase of dominance by S. tenellum prior to the establishment of either S. magellanicum or S. papillosum. Although this disturbance is of a large magnitude, it was found that equable or more severe perturbations to the water table had occurred in the past, especially at coastal sites where marine regression led to the establishment of Sphagnum-poor communities dominated by Calluna and Eriophorum vaginatum. The long- term persistence of hummock–hollow vegetation communities is demonstrated, but the appearance of persistent, more homogeneous phases suggests that this surface topography is not always well developed.