Modelling landscape effects on density-contact rate relationships of deer in eastern Alberta: Implications for chronic wasting disease

Managing wildlife diseases requires an understanding of disease transmission, which may be strongly affected by host population density and landscape features. Transmission models are typically fit from time-series disease prevalence data and modelled based on how the contact rate among hosts is affected by density, which is often assumed to be a linear (density-dependent transmission) or constant (frequency-dependent transmission) relationship. However, long-term time-series data is unavailable for emerging diseases, and this approach cannot account for independent effects of landscape. We developed a mechanistic model based on ecological data to empirically derive the contact rate-density relationship in white-tailed and mule deer in an enzootic region of chronic wasting disease (CWD) in Alberta, Canada and to determine whether it was affected by landscape. Using data collected from aerial surveys and GPS-telemetry, we developed empirical relationships predicting deer group size, home range size, and habitat selection to iteratively simulate deer distributions across a range of densities and landscapes. We calculated a relative measure of total per-capita contact rate, which is proportional to the number of other deer contacted per individual per unit time, for each distribution as the sum of pairwise contact rates between a target deer and all other individuals. Each pairwise contact rate was estimated from an empirical relationship developed from GPS-telemetry data predicting pairwise contact rates as a function of home range overlap and landscape structure. Total per-capita contact rates increased as a saturating function of density, supporting a transmission model intermediate between density- and frequency-dependent transmission. This pattern resulted from group sizes that reached an asymptote with increasing deer density, although this relationship was mediated by tree and shrub coverage in the landscape, such that in heavily wooded areas, the contact rate saturated at much lower densities. These results suggest that CWD management based on herd reductions, which require a density-dependent contact rate to be effective, may have variable effects on disease across a single management region. The novel mechanistic approach we employed for estimating effects of density and landscape on transmission is a powerful complement to typical data-fitting approaches for modelling disease transmission.     

Effect of gyrodactylid ectoparasites on host behaviour and social network structure in guppies <Emphasis Type="Italic">Poecilia reticulata</Emphasis>

Understanding how individuals modify their social interactions in response to infectious disease is of central importance for our comprehension of how disease dynamics operate in real-world populations. Whilst a significant amount of theoretical work has modelled disease transmission using network models, we have comparatively little understanding of how infectious disease impacts on the social behaviour of individuals and how these effects scale up to the level of the population. We experimentally manipulated the parasite load of female guppies (Poecilia reticulata) and introduced fish either infected with the ectoparasites Gyrodactylus spp. (experimental) or uninfected (control) into replicated semi-natural populations of eight size-matched female guppies. We quantified the behaviour and social associations of both the introduced fish and the population fish. We found that infected experimental fish spent less time associating with the population fish than the uninfected control fish. Using information on which fish initiated shoal fission (splitting) events, our results demonstrate that the population fish actively avoided infected experimental fish. We also found that the presence of an infected individual resulted in a continued decline in social network clustering up to at least 24 h after the introduction of the infected fish, whereas in the control treatment, the clustering coefficient showed an increase at this time point. These results demonstrate that the presence of a disease has implications for both the social associations of infected individuals and for the social network structure of the population, which we predict will have consequences for infectious disease transmission.     

Predation, habitat complexity, and variation in density-dependent mortality of temperate reef fishes

Density dependence in demographic rates can strongly affect the dynamics of populations. However, the mechanisms generating density dependence (e.g., predation) are also dynamic processes and may be influenced by local conditions. Understanding the manner in which local habitat features affect the occurrence and/or strength of density dependence will increase our understanding of population dynamics in heterogeneous environments. In this study I conducted two separate field experiments to investigate how local predator density and habitat complexity affect the occurrence and form of density-dependent mortality of juvenile rockfishes (Sebastes spp.). I also used yearly censuses of rockfish populations on nearshore reefs throughout central California to evaluate mortality of juvenile rockfish at large spatial scales. Manipulations of predators (juvenile bocaccio, S. paucispinus) and prey (kelp, gopher, and black-and-yellow [KGB] rockfish, Sebastes spp.) demonstrated that increasing the density of predators altered their functional response and thus altered patterns of density dependence in mortality of their prey. At low densities of predators, the number of prey consumed per predator was a decelerating function, and mortality of prey was inversely density dependent. However, at high densities of predators, the number of prey killed per predator became an accelerating response, and prey mortality was directly density dependent. Results of field experiments and large-scale surveys both indicated that the strength of density-dependent mortality may also be affected by the structural complexity of the habitat. In small-scale field experiments, increased habitat complexity increased the strength of density-dependent mortality. However, at large scales, increasing complexity resulted in a decrease in the strength of density dependence. I suggest that these differences resulted from scale-dependent changes in the predatory response that generated mortality. Whether increased habitat complexity leads to an increase or a decrease in the strength of density-dependent mortality may depend on how specific predatory responses (e.g., functional or aggregative) are altered by habitat complexity. Overall, the findings of this study suggest that rates of demographic density dependence and the resulting dynamics of local populations may largely depend upon attributes of the local habitat.     

Movement ecology: size-specific behavioral response of an invasive snail to food availability

Immigration, emigration, migration, and redistribution describe processes that involve movement of individuals. These movements are an essential part of contemporary ecological models, and understanding how movement is affected by biotic and abiotic factors is important for effectively modeling ecological processes that depend on movement. We asked how phenotypic heterogeneity (body size) and environmental heterogeneity (food resource level) affect the movement behavior of an aquatic snail (Tarebia granifera), and whether including these phenotypic and environmental effects improves advection-diffusion models of movement. We postulated various elaborations of the basic advection diffusion model as a priori working hypotheses. To test our hypotheses we measured individual snail movements in experimental streams at high- and low-food resource treatments. Using these experimental movement data, we examined the dependency of model selection on resource level and body size using Akaike's Information Criterion (AIC). At low resources, large individuals moved faster than small individuals, producing a platykurtic movement distribution; including size dependency in the model improved model performance. In stark contrast, at high resources, individuals moved upstream together as a wave, and body size differences largely disappeared. The model selection exercise indicated that population heterogeneity is best described by the advection component of movement for this species, because the top-ranked model included size dependency in advection, but not diffusion. Also, all probable models included resource dependency. Thus population and environmental heterogeneities both influence individual movement behaviors and the population-level distribution kernels, and their interaction may drive variation in movement behaviors in terms of both advection rates and diffusion rates. A behaviorally informed modeling framework will integrate the sentient response of individuals in terms of movement and enhance our ability to accurately model ecological processes that depend on animal movement.     

Species diversity modulates predation

Predation occurs in a context defined by both prey and non-prey species. At present it is largely unknown how species diversity in general, and species that are not included in a predator's diet in particular, modify predator-prey interactions. Therefore we studied how both the density and diversity of non-prey species modified predation rates in experimental microcosms. We found that even a low density of a single nonprey species depressed the asymptote of a predator's functional response. Increases in the density and diversity of non-prey species further reduced predation rates to very low levels. Controls showed that this diversity effect was not due to the identity of any of the non-prey species. Our results establish that both the density and diversity of species outside a predator's diet can significantly weaken the strength of predator-prey interactions. These results have major implications for ecological theory on species interactions in simple vs. complex communities. We discuss our findings in terms of the relationship between diversity and stability.     

The role of images of conspecifics as visual cues in the development and behavior of larval anurans

Tadpoles can alter their behavior, morphology, and life history in response to habitat change. Although chemical signals from conspecifics or predators play an important role in tadpole habitat assessment, little is known about the role of visual cues and the extent to which tadpoles rely on their vision for intraspecific social assessment. The aim of our experiments was to determine whether larval anurans use visual images of other tadpoles as indicators of density and to analyze how, and to what extent, images of conspecifics alone affect tadpole development, growth, and behavior. To assess this, we raised both Rana sylvatica and Bufo americanus tadpoles in aquaria with either quarter- or half-mirrored walls. Both physically increased density and increased density simulated with mirrors decreased tadpole growth and developmental rates, and increased activity in Rana tadpoles. Bufo tadpoles did not significantly alter their growth and development in response to visually increased density. Only true, i.e., physically, increased density had an effect on growth and activity in Bufo tadpoles. Our data show that images of conspecifics are used as visual cues by Rana tadpoles and can induce phenotypically plastic changes in several traits. This response to visual cues is taxon-specific. An erratum to this article can be found at     

Density dependence in social behaviour: home range overlap and density interacts to affect conspecific encounter rates in a gregarious ungulate

Sociality is poorly understood in the context of population processes. We used wild, female elk (Cervus canadensis) equipped with proximity-logging radio collars (n?=?62) from Manitoba, Canada (2007–2009), to test for modifying effects of population density (two areas: 0.42 and 0.22 animals/km²) on the relationship between two measures of sociality. This included the rate at which collared individuals encountered one another per year (encounters logged as animals ranging to within 1.4 m of each other) and the extent to which animals overlapped in annual home range (proportion of shared minimum convex polygon ranges). Overlap was significantly greater in the high density area compared to that of the low, but not if we only considered individuals that directly encountered each other, implying that familiar individuals will maintain a constant degree of range overlap regardless of density. Encounter rate was nonlinearly related to home range overlap. This relationship was also density-dependent, exhibiting negative density dependence at high proportions of overlap, primarily in the high density subpopulation. Sociality, as defined by two interacting measures of behaviour—encounter rate and home range overlap—exhibits a complex nonlinear relationship; we discuss the implications of these results as they pertain to sociobiology, resource competition, and pathogen transmission.     

Associations of forest bird species richness with housing and landscape patterns across the USA.

In the United States, housing density has substantially increased in and adjacent to forests. Our goal in this study was to identify how housing density and human populations are associated with avian diversity. We compared these associations to those between landscape pattern and avian diversity, and we examined how these associations vary across the conterminous forested United States. Using data from the North American Breeding Bird Survey, the U.S. Census, and the National Land Cover Database, we focused on forest and woodland bird communities and conducted our analysis at multiple levels of model specificity, first using a coarse-thematic resolution (basic models), then using a larger number of fine-thematic resolution variables (refined models). We found that housing development was associated with forest bird species richness in all forested ecoregions of the conterminous United States. However, there were important differences among ecoregions. In the basic models, housing density accounted for < 5% of variance in avian species richness. In refined models, 85% of models included housing density and/or residential land cover as significant variables. The strongest guild response was demonstrated in the Adirondack-New England ecoregion, where 29% of variation in richness of the permanent resident guild was associated with housing density. Model improvements due to regional stratification were most pronounced for cavity nesters and short-distance migrants, suggesting that these guilds may be especially sensitive to regional processes. The varying patterns of association between avian richness and attributes associated with landscape structure suggested that landscape context was an important mediating factor affecting how biodiversity responds to landscape changes. Our analysis suggested that simple, broadly applicable, land use recommendations cannot be derived from our results. Rather, anticipating future avian response to land use intensification (or reversion to native vegetation) has to be conditioned on the current landscape context and the species group of interest. Our results show that housing density and residential land cover were significant predictors of forest bird species richness, and their prediction strengths are likely to increase as development continues.     

Can rare positive interactions become common when large carnivores consume livestock?

Livestock populations in protected areas are viewed negatively because of their interaction with native ungulates through direct competition for food resources. However, livestock and native prey can also interact indirectly through their shared predator. Indirect interactions between two prey species occur when one prey modifies either the functional or numerical responses of a shared predator. This interaction is often manifested as negative effects (apparent competition) on one or both prey species through increased predation risk. But indirect interactions can also yield positive effects on a focal prey if the shared predator modifies its functional response toward increased consumption of an abundant and higher-quality alternative prey. Such a phenomenon between two prey species is underappreciated and overlooked in nature. Positive indirect effects can be expected to occur in livestock-dominated wildlife reserves containing large carnivores. We searched for such positive effects in Acacia-Zizhypus forests of India's Gir sanctuary where livestock (Bubalus bubalis and Bos indicus) and a coexisting native prey (chital deer, Axis axis) are consumed by Asiatic lions (Panthera leo persica). Chital vigilance was higher in areas with low livestock density than in areas with high livestock density. This positive indirect effect occurred because lion predation rates on livestock were twice as great where livestock were abundant than where livestock density was low. Positive indirect interactions mediated by shared predators may be more common than generally thought with rather major consequences for ecological understanding and conservation. We encourage further studies to understand outcomes of indirect interactions on long-term predator-prey dynamics in livestock-dominated protected areas.     

Stage-specific biomass overcompensation by juveniles in response to increased adult mortality in a wild fish population

Recently developed theoretical models of stage-structured consumer-resource systems have shown that stage-specific biomass overcompensation can arise in response to increased mortality rates. We parameterized a stage-structured population model to simulate the effects of increased adult mortality caused by a pathogen outbreak in the perch (Perca fluviatilis) population of Windermere (UK) in 1976. The model predicts biomass overcompensation by juveniles in response to increased adult mortality due to a shift in food-dependent growth and reproduction rates. Considering cannibalism between life stages in the model reinforces this compensatory response due to the release from predation on juveniles at high mortality rates. These model predictions are matched by our analysis of a 60-year time series of scientific monitoring of Windermere perch, which shows that the pathogen outbreak induced a strong decrease in adult biomass and a corresponding increase in juvenile biomass. Age-specific adult fecundity and size at age were higher after than before the disease outbreak, suggesting that the pathogen-induced mortality released adult perch from competition, thereby increasing somatic and reproductive growth. Higher juvenile survival after the pathogen outbreak due to a release from cannibalism likely contributed to the observed biomass overcompensation. Our findings have general implications for predicting population- and community-level responses to increased size-selective mortality caused by exploitation or disease outbreaks.     

Evaluating orangutan census techniques using nest decay rates: implications for population estimates.

An accurate estimate for orangutan nest decay time is a crucial factor in commonly used methods for estimating orangutan population size. Decay rates are known to vary, but the decay process and, thus, the temporal and spatial variation in decay time are poorly understood. We used established line-transect methodology to survey orangutan nests in a lowland forest in East Kalimantan, Indonesia, and monitored the decay of 663 nests over 20 months. Using Markov chain analysis we calculated a decay time of 602 days, which is significantly longer than times found in other studies. Based on this, we recalculated the orangutan density estimate for a site in East Kalimantan; the resulting density is much lower than previous estimates (previous estimates were 3-8 times higher than our recalculated density). Our data suggest that short-term studies where decay times are determined using matrix mathematics may produce unreliable decay times. Our findings have implications for other parts of the orangutan range where population estimates are based on potentially unreliable nest decay rate estimates, and we recommend that for various parts of the orangutan range census estimates be reexamined. Considering the high variation in decay rates there is a need to move away from using single-number decay time estimates and, preferably, to test methods that do not rely on nest decay times as alternatives for rapid assessments of orangutan habitat for conservation in Borneo.     

Primary production and rain use efficiency across a precipitation gradient on the Mongolia Plateau

Understanding how the aboveground net primary production (ANPP) of arid and semiarid ecosystems of the world responds to variations in precipitation is crucial for assessing the impacts of climate change on terrestrial ecosystems. Rain-use efficiency (RUE) is an important measure for acquiring this understanding. However, little is known about the response pattern of RUE for the largest contiguous natural grassland region of the world, the Eurasian Steppe. Here we investigated the spatial and temporal patterns of ANPP and RUE and their key driving factors based on a long-term data set from 21 natural arid and semiarid ecosystem sites across the Inner Mongolia steppe region in northern China. Our results showed that, with increasing mean annual precipitation (MAP), (1) ANPP increased while the interannual variability of ANPP declined, (2) plant species richness increased and the relative abundance of key functional groups shifted predictably, and (3) RUE increased in space across different ecosystems but decreased with increasing annual precipitation within a given ecosystem. These results clearly indicate that the patterns of both ANPP and RUE are scale dependent, and the seemingly conflicting patterns of RUE in space vs. time suggest distinctive underlying mechanisms, involving interactions among precipitation, soil N, and biotic factors. Also, while our results supported the existence of a common maximum RUE, they also indicated that its value could be substantially increased by altering resource availability, such as adding nitrogen. Our findings have important implications for understanding and predicting ecological impacts of global climate change and for management practices in arid and semiarid ecosystems in the Inner Mongolia steppe region and beyond.     

Causes and consequences of contest outcome: aggressiveness, dominance and growth in the sheepshead swordtail, Xiphophorus birchmanni

Although our understanding of how animal personality affects fitness is incomplete, one general hypothesis is that personality traits (e.g. boldness and aggressiveness) contribute to competitive ability. If so, then under resource limitation, personality differences will generate variation in life history traits crucial to fitness, like growth. Here, we test this idea using data from same-sex dyadic interaction trials of sheepshead swordtails (Xiphophorus birchmanni). In males, there was evidence of repeatable variation across a suite of agonistic contest behaviours, while repeatable opponent effects on focal behaviour were also detected. A single vector explains 80 % of the among-individual variance in multivariate phenotype and can be viewed as aggressiveness. We also find that aggressiveness predicts dominance—the repeatable tendency to win food in competition—and dominant individuals show faster post-trial weight gain (independently of initial size). In females, a dominance hierarchy predictive of weight gain was also found, but there was no evidence of variation in aggressiveness. While size often predicts contest outcome, our results show that individuals may sometimes grow larger because they are behaviourally dominant rather than vice versa. When resources are limited, personality traits such as aggression can influence growth, life history, and fitness through impacts on resource acquisition.     

Nest desertion is not predicted by cuckoldry in the Eurasian penduline tit

Engagement in extra-pair copulations is an example of the abundant conflicting interests between males and females over reproduction. Potential benefits for females and the risk of cuckoldry for males are expected to have important implications on the evolution of parental care. However, whether parents adjust parental care in response to parentage remains unclear. In Eurasian penduline tits Remiz pendulinus, which are small polygamous songbirds, parental care is carried out either by the male or by the female. In addition, one third of clutches is deserted by both male and female. Desertion takes place during the egg-laying phase. Using genotypes of nine microsatellite loci of 443 offspring and 211 adults, we test whether extra-pair paternity predicts parental care. We expect males to be more likely to desert cuckolded broods, whereas we expect females, if they obtain benefits from having multiple sires, to be more likely to care for broods with multiple paternity. Our results suggest that parental care is not adjusted to parentage on an ecological timescale. Furthermore, we found that male attractiveness does not predict cuckoldry, and we found no evidence for indirect benefits for females (i.e., increased growth rates or heterozygosity of extra-pair offspring). We argue that male Eurasian penduline tits may not be able to assess the risk of cuckoldry; thus, a direct association with parental care is unlikely to evolve. However, timing of desertion (i.e., when to desert during the egg-laying phase) may be influenced by the risk of cuckoldry. Future work applying extensive gene sequencing and quantitative genetics is likely to further our understanding of how selection may influence the association between parentage and parental care.     

Spatial anatomy of species survival: effects of predation and climate-driven environmental variability

The majority of survival analyses focus on temporal scales. Consequently, there is a limited understanding of how species survival varies over space and, ultimately, how spatial variability in the environment affects the temporal dynamics of species abundance. Using data from the Barents Sea, we study the spatiotemporal variability of the juvenile Atlantic cod (Gadus morhua) survival. We develop an index of spatial survival based on changes of juvenile cod distribution through their first winter of life (from age-0 to age-1) and study its variability in relation to biotic and abiotic factors. Over the 25 years analyzed (1980-2004), we found that, once the effect of passive drift due to dominant currents is accounted for, the area where age-0 cod survival was lowest coincided with the area of highest abundance of older cod. Within this critical region, the survival of age-0 cod was negatively affected by its own abundance, by that of older cod, and by bottom depth. Furthermore, during cold years, age-0 cod survival increased in the eastern and coldest portion of the examined area, which was typically avoided by older conspecifics. Based on these results we propose that within the examined area top-down mechanisms and predation-driven density dependence can strongly affect the spatial pattern of age-0 cod survival. Climate-related variables can also influence the spatial survival of age-0 cod by affecting their distribution and that of their predators. Results from these and similar studies, focusing on the spatial variability of survival rates, can be used to characterize species habitat quality of marine renewable resources.     

Transient population dynamics in periodic matrix models: methodology and effects of cyclic permutations

Many biological populations are subject to periodically changing environments such as years with or without fire, or rotation of crop types. The dynamics and management options for such populations are frequently investigated using periodic matrix models. However the analysis is usually limited to long-term results (asymptotic population growth rate and its sensitivity to perturbations of vital rates). In non-periodic matrix models it has been shown that long-term results may be misleading as populations are rarely in their stable structure. We therefore develop methods to analyze transient dynamics of periodic matrix models. In particular, we show how to calculate the effects of perturbations on population size within and at the end of environmental cycles. Using a model of a weed population subject to a crop rotation, we show that different cyclic permutations produce different patterns of sensitivity of population size and different population sizes. By examining how the starting environment interacts with the initial conditions, we explain how different patterns arise. Such understanding is critical to developing effective management and monitoring strategies for populations subject to periodically recurring environments.     

Insect herbivores, density dependence, and the performance of the perennial herb Solanum carolinense

How insect herbivores affect plant performance is of central importance to basic and applied ecology. A full understanding of herbivore effects on plant performance requires understanding interactions (if any) of herbivore effects with plant density and size because these interactions will be critical for determining how herbivores influence plant population size. However, few studies have considered these interactions, particularly over a wide enough range of densities to detect nonlinear effects. Here we ask whether plant density and herbivores influence plant performance linearly or nonlinearly, how plant density affects herbivore damage, and how herbivores alter density dependence in transitions between plant size classes. In a large field experiment, we manipulated the density of the herbaceous perennial plant Solanum carolinense and herbivore presence in a fully crossed design. We measured plant size, sexual reproduction, and damage to plants in two consecutive years, and asexual reproduction of new stems in the second year, allowing us to characterize both plant performance and rates of transition between plant size classes across years. We found nonlinear effects of plant density on damage. Damage by herbivores and plant density both influenced sexual and asexual reproduction of S. carolinense; these effects were mostly mediated via effects on plant size. Importantly, we found that herbivores altered the pattern of linear density dependence in some transition rates (including survival and asexual reproduction) between plant size classes. These results suggest that understanding the ecological or evolutionary effects of herbivores on plant populations requires consideration of plant density and plant size, because feedbacks between density, herbivores, and plant size may complicate longer-term dynamics.     

Interaction networks in coastal soft-sediments highlight the potential for change in ecological resilience

Recent studies emphasize the role of indirect relationships and feedback loops in maintaining ecosystem resilience. Environmental changes that impact on the organisms involved in these processes have the potential to initiate threshold responses and fundamentally shift the interactions within an ecosystem. However, empirical studies are hindered by the difficulty of designing appropriate manipulative experiments to capture this complexity. Here we employ structural equation modeling to define and test the architecture of ecosystem interaction networks. Using survey data from 19 estuaries we investigate the interactions between biological (abundance of large bioturbating macrofauna, microphytobenthos, and detrital matter) and physical (sediment grain size) processes. We assess the potential for abrupt changes in the architecture of the network and the strength of interactions to occur across environmental gradients. Our analysis identified a potential threshold in the relationship between sediment mud content and benthic chlorophyll a, at -12 microg/g, using quantile regression. Below this threshold, the interaction network involved different variables and fewer feedbacks than above. This approach has potential to improve our empirical understanding of thresholds in ecological systems and our ability to design manipulative experiments that test how and when a threshold will be passed. It can also be used to indicate to resource managers that a particular system has the potential to exhibit threshold responses to environmental change, emphasizing precautionary management and facilitating a better understanding of how persistent multiple stressors threaten the resilience and long-term use of natural ecosystems.     

Behavioral responses of territorial red squirrels to natural and experimental variation in population density

The relative scarcity of studies at the intersection of behavioral and population ecology is surprising given the presumed importance of behavior in density-dependent population regulation. Here we tested whether North American red squirrels (Tamiasciurus hudsonicus) adjust their behavior in response to local population density and whether they use rates of territorial vocalizations in their local neighborhood to assess density. We examined these relationships using 18?years of live trapping and 20?years of behavioral data that were collected across natural variation in local population density. To disentangle the effects of population density on behavior from those due to changes in per capita food abundance or changes in the frequency of antagonistic interactions with neighbors, we also experimentally manipulated population density with long-term food supplementation as well as perceived population density with long-term playbacks of territorial vocalizations. The frequency with which squirrels emitted territorial vocalizations was positively associated with local population density. In contrast, antagonistic physical interactions observed between squirrels and territorial intrusions were rare and the frequency of intrusions was weakly and negatively, not positively, associated with population density. Squirrels experiencing naturally and experimentally high density conditions spent less time in the nest and feeding but more time being vigilant. Similar density-dependent changes in behavior were observed in response to our manipulations of perceived population density, indicating that vocalization rates and not physical interactions or food abundance were the mechanism by which squirrels assessed and responded behaviorally to changes in local density.     

Intraspecific competition in<Emphasis Type="Italic"> Fucus serratus</Emphasis> and<Emphasis Type="Italic"> F</Emphasis>.<Emphasis Type="Italic"> evanescens</Emphasis> (Phaeophyceae: Fucales) germlings: effects of settlement density,nutrient concentration,and temperature

Survival and growth of early post-settlement stages are critical for the development of seaweed populations. Fucoid germlings commonly settle in dense monospecific aggregates, where intraspecific competition and environmental variables (e.g. nutrient concentration and temperature) may affect survival and growth. Using factorial experiments, we determined the effects of settlement density (~10, ~50 and ~250 germlings cm^–2), nutrient enrichment (from ~10 to ~40 µM N and from ~0.5 to ~2.5 µM P), and temperature (7°C and 17°C) on Fucus serratus and F. evanescens germlings in laboratory cultures over 3 months. Settlement density, nutrient concentration and temperature interactively affected growth of germlings, and the magnitude of this interaction varied between the two species. This represents the first record of such factorial interactions in Fucus spp. germlings. Intraspecific competition, estimated as the relative reduction in germling growth and survival from low to high densities, increased with decreasing nutrient concentration and increasing temperature in both species. While temperature and nutrient concentration had little effect on germling size distributions, size inequality and skewness generally increased with germling density, indicating that a few large individuals gained dominance and suppressed many smaller ones at high density. Self-thinning increased with settlement density and depended on nutrient concentration and species at high density. At high density, self-thinning increased with decreasing nutrient levels in F. evanescens, but not in F. serratus. At low density, nutrient enrichment increased germling growth in F. evanescens, but not in F. serratus, whereas growth in both species was stimulated by nutrient enrichment at higher densities. These results suggest that germling growth and self-thinning are more sensitive to variation in nutrient concentration in F. evanescens than in F. serratus. The potential implications of our findings for the understanding of eutrophication-related abundance changes in both species in southern Norway are discussed.Communicated by L. Hagerman, Helsingør