Inferring controls on the epidemiology of beech bark disease from spatial patterning of disease organisms

• 1 Spatial pattern in the distribution and abundance of organisms is an emergent property of collective rates of reproduction, survival and movement of individuals in a heterogeneous environment.
• 2 The form, intensity and scale of spatial patterning can be used to test hypotheses regarding the relative importance of candidate processes to population dynamics.
• 3 Using 84 plots across eastern North America, we studied populations of two associated plant parasites, the invasive felted beech scale Cryptococcus fagisuga Lind. and the native Neonectria fungi, which together cause beech bark disease (BBD).
• 4 We evaluated spatial patterns at the scales of trees within stands, stands within the forest and forests within the landscape to examine four hypothetically important factors in the ecology of the disease: (i) local contagion within stands; (ii) regional contagion, or among patch infection–reinfection dynamics; (iii) variation in host susceptibility linked to genetic and/or environmental heterogeneity; and (iv) climate effects on population growth of BBD organisms.
• 5 Analyses revealed an unexpected lack of spatial aggregation in BBD populations among trees, stands and forests. This implies that propagule pressure is generally sufficiently high throughout the infested region of North America such that neither trees nor stands are spared from the disease by dispersal limitations of the disease agents. Furthermore, variation in tree and stand level susceptibility has minimal impact on BBD dynamics and climate is not a conspicuous driver of abundance within the core range of BBD.

     

Vertical stratification in the spatial distribution of the beech scale insect (Ultracoelostoma assimile) in Nothofagus tree canopies in New Zealand

Abstract. 1. The degree of infestation by New Zealand sooty beech scale insects (Ultracoelostoma assimile, Homoptera: Margarodidae) varies dramatically among adjacent southern beech trees (Nothofagus spp., Fagaceae), but has previously been assumed to be uniformly or randomly distributed within individual host trees. In this study, a full‐census survey was conducted from ground level to canopy level on 14 naturally occurring, canopy‐dominant red beech (Nothofagus fusca) trees (size range 38.7–107.6 cm diameter at breast height) to determine the degree of within‐tree heterogeneity in herbivore density. 2. The within‐tree distribution of the sooty beech scale was vertically stratified and highly heterogeneous, with the greatest densities occurring on bark surfaces in the canopy rather than on the trunk, and on the lower rather than upper sides of the branches. The spatial distribution was strongly negatively correlated with trunk and branch diameter, and increasing bark thickness (as a function of diameter) provides a plausible explanation for differences in the establishment and population density of sooty beech scale insects with trunk and branch size. Furthermore, there was a significant change in the spatial distribution of scale insect populations on trunks and branches of trees of increasing diameter at breast height. This indicates a strong temporal component to the spatial dynamics of the sooty beech scale insect driven by changing host phenology. Future studies on phytophagous insects infesting large host trees need to consider more explicitly changes in population dynamics through space and time. 3. Because of the high degree of within‐tree heterogeneity in population density, the total population size of scale insects on an individual tree could not be predicted from any measure of population density low on the trunk. However, the dry weight biomass of sooty mould fungi growing on the ground beneath infested trees was a remarkably accurate predictor of the total population size of scale insects. The use of sooty mould fungi as a relative measure of population size could be incorporated into studies of other honeydew‐producing hemipterans, since the growth of sooty mould is a distinctive feature synonymous with high concentrations of honeydew production worldwide.     

Population dynamics of the beech caterpillar, <Emphasis Type="Italic">Syntypistis punctatella</Emphasis>, and biotic and abiotic factors

The beech caterpillar, Syntypistis punctatella (Motschulsky) (Lepidoptera: Notodontidae), often causes extensive defoliation of beech forests in Japan. Outbreaks have often occurred synchronously among different areas at intervals of 8–11 years. Synchrony of outbreaks was considered to be caused by synchrony of weather. Populations of this insect exhibit periodical dynamics in both outbreak and nonoutbreak areas. Factors that might influence the population dynamics of the beech caterpillar were classified from the point of view of the natural bioregulation com-plex, which includes a coleopteran predator, Calosoma maximowiczi, avian predators, parasitoids, entomopathogenic fungi, and delayed induced defensive response (DIR) of beech trees. Because such periodic population dynamics are believed to be caused by one or more delayed density-dependent factors, delayed density-dependent mortality has been identified as a likely source of population cycles. The DIR and pathogenic diseases showed a high order of density dependence. An infectious pathogen, Cordyceps militaris, was considered to be the most plausible agent responsible for periodic dynamics of the beech caterpillar population because insect diseases were effective in cases in which the S. punctatella population started to decrease without reaching outbreak densities, but DIR was not. Conspicuous defoliation caused by this insect tends to occur at certain elevations, where forests are composed of pure stands of beech trees. I propose three different hypotheses to explain this phenomenon: the diversity–stability hypothesis, the resource concentration hypothesis, and the altitudinal soil nutrient hypothesis. Received: November 20, 1999 / Accepted: August 3, 2000     

The effects of mice on stoats in southern beech forests

Introduced stoats (Mustela erminea) are important invasive predators in southern beech (Nothofagus sp.) forests in New Zealand. In these forests, one of their primary prey species – introduced house mice (Mus musculus), fluctuate dramatically between years, driven by the irregular heavy seed‐fall (masting) of the beech trees. We examined the effects of mice on stoats in this system by comparing the weights, age structure and population densities of stoats caught on two large islands in Fiordland, New Zealand – one that has mice (Resolution Island) and one that does not (Secretary Island). On Resolution Island, the stoat population showed a history of recruitment spikes and troughs linked to beech masting, whereas the Secretary Island population had more constant recruitment, indicating that rodents are probably the primary cause for the ‘boom and bust’ population cycle of stoats in beech forests. Resolutions Island stoats were 10% heavier on average than Secretary Island stoats, supporting the hypothesis that the availability of larger prey (mice verses wētā) leads to larger stoats. Beech masting years on this island were also correlated with a higher weight for stoats born in the year of the masting event. The detailed demographic information on the stoat populations of these two islands supports previously suggested interactions among mice, stoats and beech masting. These interactions may have important consequences for the endemic species that interact with fluctuating populations of mice and stoats.     

Differing contributions of density dependence and climate to the population dynamics of three eruptive herbivores

1. Although both endogenous and exogenous processes regulate populations, the current understanding of the contributions from density dependence and climate to the population dynamics of eruptive herbivores remains limited. 2. Using a 17‐year time series of three cereal aphid species [Rhopalosiphum padi L., Metopolophium dirhodum (Walker), and Diuraphis noxia (Kurdumov)] compiled from a trapping network spanning the northwestern U.S.A., temporal and spatial patterns associated with population fluctuations, and modelled density dependence in aphid abundances were tested. These models were used to analyse correlations between climate and aphid abundances in the presence and absence of residual variance as a result of density‐dependent effects. 3. The temporal dynamics of aphid population fluctuations indicated periodicity, with no clear evidence for a spatial pattern underlying population fluctuations. 4. Aphid abundances oscillated in a manner consistent with delayed density dependence for all three aphid species, although the strength of these feedbacks differed among species. 5. Diuraphis noxia abundances were negatively correlated with increasing temperatures in the absence of density‐dependent effects, whereas M. dirhodum abundances were positively correlated with increasing cumulative precipitation in the presence of density‐dependent effects; yet, R. padi abundances were unrelated to climate variables irrespective of population feedbacks. 6. Our analysis suggests that endogenous feedbacks differentially regulate aphid populations in the northwestern U.S.A., and these feedbacks may operate at an expansive spatial scale. It is concluded that the contributions of density dependence and climate to aphid population dynamics are species‐specific in spite of similar ecological niches, with implications for assessing species responses to climate variability.     

Early-stage of invasion by beech bark disease does not necessarily trigger American beech root sucker establishment in hardwood stands

Two concomitant phenomena currently affect the dynamics of sugar maple-American beech (AB) stands in northeastern North America: beech bark disease (BBD), and increased AB understory density. Many studies suggest a causal link between the two phenomena, i.e., BBD favouring beech regeneration. But this link has yet to be experimentally demonstrated. To address the question, we compared regeneration composition between recently BBD-affected and -unaffected stands. A total of 109 stands were sampled; half were affected by BBD. Seedling and sapling density were assessed, together with the origin (seedling or sprout). While BBD affects stands in the eastern part of the study region, AB was observed in the understory across the entire region. No clear difference in AB sprout density between BBD-affected and -unaffected stands was observed while AB seedling density—as well as pooled AB seedling and sprout density were higher in unaffected stands. Findings suggests that BBD, in its early stage, is not a necessary trigger of AB understory establishment. Yet, AB sapling basal area generally was higher in stands affected by BBD, likely indicating a greater rate of AB understory development due to increased light availability beneath a more open crown canopy. That development can lead to AB understory dominance. This distinction—BBD not necessarily triggering AB root sucker establishment but favoring AB advance regeneration development—also questions the generalized perception that dense AB thickets necessarily originate from root suckers.     

Spatially patterned guild structure in larval parasitoids of cyclically outbreaking winter moth populations

1. Wide temporal fluctuations in host abundance are a potential source of instability and stochasticity in the spatiotemporal population dynamics of associated parasitoid species. Within parasitoid guilds (i.e. parasitoids with similar modes of host utilisation), a conceivable outcome is guild organisation according to a lottery model, in which guild members attain local dominance by colonising previously emptied habitats during increasing host density, before other guild members. In the spatial dimension, an expected manifestation of such dynamics is variable guild structure even across homogeneous habitats. 2. We examined the extent of large‐scale spatial patterning of guild characteristics in larval parasitoid wasps associated with cyclically outbreaking populations of the geometrid moth Operophtera brumata in northern Fennoscandia. The study was performed at the onset of the crash‐phase of the geometrid's outbreak cycle, along a 70‐km transect in costal northern Norway, characterised by largely homogeneous environmental conditions, except for a small climatic gradient. 3. There was a distinct large‐scale spatial turnover in dominance among the major parasitoid groups (i.e. guild structure) in O. brumata along the transect, whereas the total prevalence rate of the guild and its diversity showed no consistent variation. Guild structure was unrelated to host density. 4. Although group‐specific responses to a slight spatial climatic gradient cannot be rejected as a causal mechanism, we conclude that our results are consistent with the expectation from large‐scale stochastic extinction‐recolonisation dynamics among functionally equivalent parasitoids relying on a host with strongly cyclic population fluctuations.     

Adult trees cause density‐dependent mortality in conspecific seedlings by regulating the frequency of pathogenic soil fungi

Negative density‐dependent seedling mortality has been widely detected in tropical, subtropical and temperate forests, with soil pathogens as a major driver. Here we investigated how host density affects the composition of soil pathogen communities and consequently influences the strength of plant‐soil feedbacks. In field censuses of six 1‐ha permanent plots, we found that survival was much lower for newly germinated seedlings that were surrounded by more conspecific adults. The relative abundance of pathogenic fungi in soil increased with increasing conspecific tree density for five of nine tree species; more soil pathogens accumulated around roots where adult tree density was higher, and this greater pathogen frequency was associated with lower seedling survival. Our findings show how tree density influences populations of soil pathogens, which creates plant‐soil feedbacks that contribute to community‐level and population‐level compensatory trends in seedling survival.     

Ectomycorrhizal fungal diversity decreases in Mediterranean pine forests adapted to recurrent fires

Fire is a major disturbance linked to the evolutionary history and climate of Mediterranean ecosystems, where the vegetation has evolved fire‐adaptive traits (e.g., serotiny in pines). In Mediterranean forests, mutualistic feedbacks between trees and ectomycorrhizal (ECM) fungi, essential for ecosystem dynamics, might be shaped by recurrent fires. We tested how the structure and function of ECM fungal communities of Pinus pinaster and Pinus halepensis vary among populations subjected to high and low fire recurrence in Mediterranean ecosystems, and analysed the relative contribution of environmental (climate, soil properties) and tree‐mediated (serotiny) factors. For both pines, local and regional ECM fungal diversity were lower in areas of high than low fire recurrence, although certain fungal species were favoured in the former. A general decline of ECM root‐tip enzymatic activity for P. pinaster was associated with high fire recurrence, but not for P. halepensis. Fire recurrence and fire‐related factors such as climate, soil properties or tree phenotype explained these results. In addition to the main influence of climate, the tree fire‐adaptive trait serotiny recovered a great portion of the variation in structure and function of ECM fungal communities associated with fire recurrence. Edaphic conditions (especially pH, tightly linked to bedrock type) were an important driver shaping ECM fungal communities, but mainly at the local scale and probably independently of the fire recurrence. Our results show that ECM fungal community shifts are associated with fire recurrence in fire‐prone dry Mediterranean forests, and reveal complex feedbacks among trees, mutualistic fungi and the surrounding environment in these ecosystems.     

Long-term Changes in Forest Carbon and Nitrogen Cycling Caused by an Introduced Pest/Pathogen Complex

Invasion of exotic forest pests and pathogens is a serious environmental problem for many forests throughout the world, and has been especially damaging to forests of eastern North America. We studied the impacts of an exotic pest/pathogen complex, the beech bark disease (BBD), in the Catskill Mountains of New York State, USA. In this region, BBD has caused a decline in the basal area of American beech (Fagus grandifolia Ehrh.) over the last 60 years and this decline has been accompanied by an increase in the basal area of sugar maple (Acer saccharum Marsh.). We studied the impacts of the BBD on carbon (C) and nitrogen (N) cycling using a series of stands that represented a sequence of disease impact and beech replacement by sugar maple. Our study showed that these long-term changes in tree species composition can lead to important changes in C and N cycling in the ecosystem, including an increase in litter decomposition, a decrease in soil C:N ratio, and an increase in extractable nitrate in the soil and nitrate in soil solution. Rates of potential net N mineralization and nitrification did not change across the BBD sequence, but the fraction of mineralized N that was nitrified increased significantly. Many of the observed changes in ecosystem function are larger in magnitude than those attributed to climate change or air pollution, suggesting that the impacts of invasive pests and pathogens on tree species composition could be one of the most important factors driving changes in C and N cycling in these forests in the coming decades.     

Density-dependent effects on the reproductive fitness of the New Zealand beech scale insect (Ultracoelostoma assimile) across multiple spatial scales

Abstract.  1. Intraspecific competition can be an important factor influencing the individual fitness of organisms. This study reports density-dependent effects on the fecundity of the beech scale insect ( Ultracoelostoma assimile , Hemiptera: Margarodidae) on naturally occurring, canopy-dominant red beech ( Nothofagus fusca , Fagaceae) trees in New Zealand. For the first time an increasing intensity of intraspecific competition at increasing spatial scales within individual host trees is demonstrated.
2. Beech scale insect 'tests' containing adult female scale insects and eggs were collected from the trunks of 10 red beech trees [17.7–48.5 cm in diameter at breast height (1.4 m), d.b.h.] with varying densities of scale insect infestation. The relationship between individual female fecundity and scale insect density at three spatial scales: local (within 5 cm), lower trunk (below 2 m above ground level), and whole tree, was tested.
3. Beech scale insect fecundity was density dependent, with total female egg load decreasing with increasing scale insect density. The strength of the density-dependent effect increased with increasing spatial scale, suggesting that scale insects are creating a tree-wide drain on the quality of phloem sap, rather than depleting nutrients from localised, high-density areas within trees. These results indicate that at high densities, the New Zealand beech scale insect can have a negative effect on the nutritional quality of the phloem of red beech, thus negatively affecting conspecifics elsewhere on individual host trees. The increasing intensity of the effect with increasing spatial scale within individual trees emphasises the importance of measuring density-dependent effects at the appropriate spatial scale.     

Population dynamics of insects associated with Rumex obtusifolius in different habitats

The effect of different habitats on insect communities associated with Rumex obtusifolius was studied in detail. Study sites were selected for their variety of environmental characteristics (vegetation and habitat): shaded woodland, disturbed wasteland and pasture. The results of this study were that insect populations associated with R. obtusifolius responded differently to the different habitats. The observed differences in population dynamics among the insects showed quite clearly their responses to different habitats. There was also a pattern of time partitioning in the exploitation of the host‐plant among the insect species observed. Philaenus spumarius at nymphal stage had a peak density on approximately 20 June, Apion frumentarium and Apion spp. (Apion species other than A. frumentarium) had a peak density on approximately 30 June and Aphis rumicis had a peak density on approximately 28 July. This pattern of shift in time for peak population density (time‐shift) among the insects sharing a common host‐plant may guarantee each insect the opportunity to exploit the plant to a maximum.     

Spatial Simulation of Epizootics Caused by Beauveria bassiana in Russian Wheat Aphid Populations

Models of disease dynamics commonly make the assumption of spatial homogeneity in the underlying host population. However, insect behavior may result in spatially heterogeneous populations with which pathogens interact. We modified a simulation model of temporal and spatial population dynamics of the Russian wheat aphid, Diuraphis noxia, on preferred or nonpreferred host plants, by incorporating effects of the entomopathogenic fungus, Beauveria bassiana. Epizootic parameters included time from inoculation of aphids until death, duration of sporulation, and estimated exposure probability. Simulations first predicted results of previously described experiments in which D. noxia adults were inoculated with conidial suspensions or water and placed on wheat or oat seedlings in 81-plant grids in cages. Subsequently, large-scale simulations were run for hypothetical field situations on 50 × 50-plant grids of wheat or oat. With B. bassiana present for both cage and larger scale simulations, results indicated that, on oat, an expanding infection front lagged behind the expanding aphid population front. Continual aphid movement from hosts resulted in many escapes, and the aphid population persisted at slightly reduced levels. On the preferred wheat host, patterns developed with pockets of infected aphids and other pockets of healthy aphids. Localized aphid populations that escaped initial infestation were able to proliferate, whereas other local populations were greatly reduced or became extinct due to lack of movement from the hosts, resulting in increased exposure to pathogen inoculum. Thus, proliferation and fluctuation of the pathogen were strongly influenced by the plant hosts' effects on aphid movement behavior. Incorporating spatial dynamics into disease models should prove useful in other efforts to predict biological control efficacy by entomopathogenic fungi in heterogeneous habitats.     

Effects of added nitrogen on leaf nitrogen concentration and population dynamics of insects associated with Rumex obtusifolius L.

A series of experiments was designed to study the effects of changes in leaf nitrogen (N) concentration on population dynamics of a range of insects associated with Rumex obtusifolius. During the study period, the N concentration in leaves fluctuated between 2 and 6% dry weight. The insects at both study sites showed a pattern of shift in time for peak population density (time‐shift). This suggests that the insect community dynamics might be closely related to the allocation and distribution of resources by the host‐plant. The elevated N concentration in the host‐plant did not change the temporal order of the peak population densities of the insects, although it may determine the time for the optimal exploitation of resources by each insect species.     

Complex fine‐scale phylogeographical patterns in a putative refugial region for Fagus sylvatica (Fagaceae)

Broad‐scale plastid (chloroplast) DNA studies of beech (Fagus sylvatica) populations suggest the existence of glacial refugia and introgression zones in south‐eastern Europe. We choose a possible refugium of beech in northern Greece, Mt. Paggeo, which hosts a private plastid haplotype for beech, to conduct a fine‐scale genetic study. We attempt to confirm or reject the hypothesis of the existence of a small‐scale refugium and to gain an understanding of the ecological and topographical factors affecting the spatial distribution of plastid haplotypes in the area. Our results reveal a high haplotype diversity on Mt. Paggeo, but the overall distribution of haplotypes shows no significant correlation with the ecological characteristics of the beech forests. However, the private haplotype is found at high frequencies in beech forests located in or near ravines, having a high spatial overlap with a relict vegetation type occurring in ecological conditions found mainly in ravines. This result emphasizes the importance of topography in the existence of glacial refugia in the wider area. Furthermore, haplotypes originating from two more widespread beech lineages in Greece are found on Mt. Paggeo, indicating a possible mixing of populations originating from a local refugium with populations from remote refugia that possibly migrated into the area after the last glaciation. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 516–528.     

Differential Responses of Herbivores and Herbivory to Management in Temperate European Beech

Forest management not only affects biodiversity but also might alter ecosystem processes mediated by the organisms, i.e. herbivory the removal of plant biomass by plant-eating insects and other arthropod groups. Aiming at revealing general relationships between forest management and herbivory we investigated aboveground arthropod herbivory in 105 plots dominated by European beech in three different regions in Germany in the sun-exposed canopy of mature beech trees and on beech saplings in the understorey. We separately assessed damage by different guilds of herbivores, i.e. chewing, sucking and scraping herbivores, gall-forming insects and mites, and leaf-mining insects. We asked whether herbivory differs among different forest management regimes (unmanaged, uneven-aged managed, even-aged managed) and among age-classes within even-aged forests. We further tested for consistency of relationships between regions, strata and herbivore guilds. On average, almost 80% of beech leaves showed herbivory damage, and about 6% of leaf area was consumed. Chewing damage was most common, whereas leaf sucking and scraping damage were very rare. Damage was generally greater in the canopy than in the understorey, in particular for chewing and scraping damage, and the occurrence of mines. There was little difference in herbivory among differently managed forests and the effects of management on damage differed among regions, strata and damage types. Covariates such as wood volume, tree density and plant diversity weakly influenced herbivory, and effects differed between herbivory types. We conclude that despite of the relatively low number of species attacking beech; arthropod herbivory on beech is generally high. We further conclude that responses of herbivory to forest management are multifaceted and environmental factors such as forest structure variables affecting in particular microclimatic conditions are more likely to explain the variability in herbivory among beech forest plots.     

Species richness,abundance and diversity of ichneumonid wasps in Japanese beech forests impacted by sika deer and sawfly herbivory

We investigated the community structure of ichneumonid wasps inhabiting beech forests at six sites in the Tanzawa Mountains of Japan under different magnitudes of impact by sika deer Cervus nippon and beech sawfly Fagineura crenativora on vegetation. Using yellow flight-interception traps, we captured 2,528 ichneumonid wasps representing 367 species in 23 subfamilies. The number of species at each site ranged from 77 to 136 and approximately 80% of these were low-density species (i.e. only one to two individuals captured per site). The number of individuals at each site ranged from 248 to 897, and the percentage of the beech sawfly parasitoids varied widely from 1% to 57%. The numbers of species in parasitoid groups categorized according to their hosts, that is, sawfly (not including the beech sawfly), Lepidoptera, woodborer, fungivore or the others, did not greatly differ among the study sites. Parasitoids attacking herbivorous insects exceeded others in species richness and abundance at all sites. Six sites were classified into four groups in terms of abundance of the host groups when excluding the parasitoids of the beech sawfly, but into only two groups when including these parasitoids. Species diversity and evenness were the highest at the least impacted site even if the beech sawfly parasitoids were excluded from calculation. We suggested some environmental factors, such as groundcover vegetation, abundance of the beech sawfly and structure and age of forest stands, that could have affected the community structure of ichneumonid wasps in the beech forests.     

病原菌与自然植物种群Ⅱ病原菌与植物种群生物学

病原菌在自然植物种群中普遍存在,其对寄主植物的生长发育,对寄主植物种群的大小、结构、动态、遗传和进化等都有重要影响。本文着重论述：１）病原菌对寄主植物个体的影响;２）病原菌对寄主植物种群生物学的影响;３）菌病发生的空间格局;４）病原菌感染的种群模型。     

Whether larval amphibians school does not affect the parasite aggregation rule: testing the effects of host spatial heterogeneity in field and experimental studies

Almost all macroparasites show over‐dispersed infections within natural host populations such that most parasites are distributed among a few heavily‐infected individuals. Despite the importance of parasite aggregation for understanding system stability, the potential for population regulation, and super‐spreading events, many questions persist about its underlying drivers. Theoretically, aggregation results from heterogeneity in host exposure, resistance, and tolerance. However, few studies have examined how host spatial arrangement – which likely affects both parasite encounter and density‐dependent interactions – influences infection and dispersion, representing a critical gap in our current knowledge regarding the possible drivers of parasite aggregation. Using field data from over 165 ponds and 8000 hosts, we evaluated how the spatial clustering of amphibian larvae within ponds 1) varied among different amphibian species, and 2), affected the distribution of parasites within the host population using Taylor's power law. A complementary mesocosm experiment used field‐guided manipulations of the spatial arrangement of larval amphibians to create a gradient in host clustering while controlling host density, thereby testing for spatial effects on both infection success and aggregation by three different trematode species. Our field data indicated that larval amphibians exhibited significant spatial clustering that was well captured by Taylor's power law (R² 0.92 to 0.97 for different host species), but the residual variation only weakly correlated with observed patterns of trematode parasite over‐dispersion. Correspondingly, experimental manipulation of host clustering had no effects on parasite infection success or the degree of parasite aggregation among cages or mesocosms. Given the importance of parasite over‐dispersion for host populations and disease dynamics, we advocate for further investigations of host and parasite spatial aggregation, particularly studies that incorporate and/or control for heterogeneity in exposure and susceptibility.     

Nutrient enrichment alters dynamics in experimental plant populations

Controlling weed populations requires an understanding of their underlying population dynamics which can be achieved through a combination of model development and long-term studies. In this paper, we develop models based on long-term data from experimental populations of the weedy annual plant Cardamine pensylvanica. Four replicate populations of C. pensylvanica were grown in growth chambers under three different nutrient levels but with all other environmental conditions held constant. We analyze the resulting time series using generalized additive models and perform stability analyses using Lyapunov exponents. Further, we test whether the proposed mechanism, delayed density dependence caused by maternal effects, is operating in our system by experimentally manipulating maternal density and assessing the resulting offspring quality. Our results show that that increasing the frequency of nutrients causes plant population dynamics to shift from stable to damped 2-point oscillations to longer cycles. This shift in population dynamics is due to a shift at high nutrients from populations being regulated by first order density feedbacks to being regulated by both first order and second order density feedbacks. A consequence of these first order and second order feedbacks was an increase in cycle lengths as demonstrated by the presence of complex eigenvalues. A short-term experiment confirmed that when grown under high nutrients, the density of maternal plants strongly affected offspring size, providing a mechanism whereby these second order density feedbacks could operate. Our results demonstrate that increasing nutrient frequency results in a qualitative shift in dynamics from stable to longer cycles.