Analysis of the relationship between allozyme heterozygosity and fitness in the rare Gentiana pneumonanthe L.

Especially for rare species occurring in small populations, which are prone to loss of genetic variation and inbreeding, detailed knowledge of the relationship between heterozygosity and fitness is generally lacking. After reporting on allozyme variation and fitness in relation to population size in the rare plant Gentiana pneumonanthe, we present a more detailed analysis of the association between heterozygosity and individual fitness. The aim of this study was to test whether increased fitness of more heterozygous individuals is explained best by the ‘inbreeding’ hypothesis or by the ‘overdominance’ hypothesis. Individual fitness was measured during 8 months of growth in the greenhouse as the performance for six life-history parameters. PCA reduced these parameters to four main Fitness Components. Individual heterozygosity was scored for seven polymorphic allozyme loci. For some of these loci (e.g. Aat3, Pgm1 and 6Pgdh2) heterozygotes showed a significantly higher relative fitness than homozygotes. To test the inbreeding model, regression analyses were performed between each Fitness Component and the number of heterozygous loci per individual. Multiple regressions with the adaptive distance of five loci as independent variables were used to test the overdominance model. Only the inbreeding model was a statistically significant explanation for the relationship between heterozygosity and fitness in G. pneumonanthe. The number of heterozygous loci was significantly negatively correlated with the coefficients of variation of three of the six initially measured fitness parameters. This suggests a lower developmental stability among more homozygous plants and may explain the higher phenotypic variation in small populations of the species observed earlier. The importance of the results for conservation biology is discussed.     

The effects of fire and fragmentation on occurrence and flowering of a rare perennial plant

The pine rocklands of southern Florida are a fire-dependent forest associated with outcroppings of limestone. Pine rockland plants have several adaptations to fire, and for many species burns increase plant growth, flowering, and seedling establishment. The pine rockland forest has been reduced and fragmented in recent decades. Outside of Everglades National Park, only two percent of the original pine rocklands remain, and are in the form of small fragments. Habitat fragmentation may have a negative effect on the biology of plants; we investigated the effects of both fire and fragmentation on Angadenia berteroi (A.DC.) Miers, a threatened species of the southern Florida pine rockland. We estimated the density and flowering of A. berteroi using adaptive cluster sampling in six study sites with different fire and disturbance histories. A. berteroi is more abundant in the largest fragments, and those having experienced fire most recently. However, fragmentation and lack of fire did not appear to have a great impact on flowering or fruit production. Insights from this threatened species may provide impetus not only to conserve, but to properly manage remaining pine rocklands in south Florida.     

Temporal and spatial variation in age-specific survival rates of a long-lived mammal, the Hawaiian monk seal

Estimates of variability in pinniped survival rates are generally based on observations at single sites, so it is not certain whether observed rates represent the whole population. Here, we provide a comprehensive analysis of spatio-temporal variation in age-specific survival rates for endangered Hawaiian monk seals (Monachus schauinslandi) based on capture-recapture analyses of more than 85% of the pups weaned in this population over the last two decades. Uniquely, these data have been collected from six subpopulations, encompassing all major breeding sites across its 1800 km long core range. Analyses of individual subpopulations revealed similar patterns in age-specific survival, characterized by the relatively low survival rates from weaning to 2 years of age, intermediate rates to 4 years of age, and then by relatively high 'mature' survival rates until 17 years of age, after which a senescent decline was observed. Juvenile, subadult and adult survival rates all varied significantly over time. Trends in survival among subpopulations were coherent with their relative geographical positions, suggesting regional structuring and connectedness within the archipelago. Survival rates for different age classes tended to be positively correlated, suggesting that similar factors may influence the survival for seals of all ages.     

Influence of herbivores on a perennial plant: variation with life history stage and herbivore species

Herbivores have diverse impacts on their host plants, potentially altering survival, growth, fecundity, and other aspects of plant performance. Especially for longer-lived plant species, the effects of a single herbivore species can vary markedly throughout the life of the host plant. In addition, the effects of herbivory during any given life history stage of a host plant may also vary considerably with different types of herbivores. To investigate the effects of herbivory by black-tailed deer (Odocoileus hemionus columbianus) and snails (Helminthoglypta arrosa and Helix aspersa) on a nitrogen-fixing shrub, Lupinus chamissonis, we established three exclosure experiments in a sand dune system on the coast of northern California. These experiments documented that deer browsing significantly reduced the volume and growth rate of lupines in the seedling and juvenile life stages. Since plant volume was strongly correlated with aboveground dry biomass for lupines, such herbivore-induced reductions in volume should translate into losses of aboveground biomass. Deer browsing also significantly altered the likelihood of attack by and density of a leaf-galling cecidomyid fly (Dasineura lupinorum), suggesting that a vertebrate herbivore indirectly affected an invertebrate herbivore in this system. Although deer did not significantly affect the survival of lupine seedlings and juveniles, individuals protected from deer had consistently greater survival in the two separate experiments. Our results revealed that snails did not have a significant effect on the survival or growth of juvenile plants, despite being common on and around lupines. An exclosure experiment revealed that herbivory by deer significantly reduced the shoot lengths of mature shrubs, but led only to a minimal reduction in growth rates. In addition, we found that browsed shrubs had significantly greater inflorescence production, but also produced individual seeds with significantly reduced mass. Collectively, these data indicate that deer and snails have widely differing effects on their shared host plant; browsing by deer indirectly affects insect herbivores, and the impacts of deer change markedly with the life history stage of their host plant.     

Temporal variation of juvenile survival in a long-lived species: the role of parasites and body condition

Studies of population dynamics of long-lived species have generally focused on adult survival because population growth should be most sensitive to this parameter. However, actual variations in population size can often be driven by other demographic parameters, such as juvenile survival, when they show high temporal variability. We used capture–recapture data from a long-term study of a hunted, migratory species, the greater snow goose (Chen caerulescens atlantica), to assess temporal variability in first-year survival and the relative importance of natural and hunting mortality. We also conducted a parasite-removal experiment to determine the effect of internal parasites and body condition on temporal variation in juvenile survival. We found that juvenile survival showed a higher temporal variability than adult survival and that natural mortality was more important than hunting mortality, unlike in adults. Parasite removal increased first-year survival and reduced its annual variability in females only. Body condition at fledging was also positively correlated with first-year survival in treated females. With reduced parasite load, females, which are thought to invest more in their immune system than males according to Bateman’s principle, could probably reallocate more energy to growth than males, leading to a higher survival. Treated birds also had a higher survival than control ones during their second year, suggesting a developmental effect that manifested later in life. Our study shows that natural factors such as internal parasites may be a major source of variation in juvenile survival of a long-lived, migratory bird, which has implications for its population dynamics.     

The demographic impact of extreme events: stochastic weather drives survival and population dynamics in a long-lived seabird

1. Most scenarios for future climate change predict increased variability and thus increased frequency of extreme weather events. To predict impacts of climate change on wild populations, we need to understand whether this translates into increased variability in demographic parameters, which would lead to reduced population growth rates even without a change in mean parameter values. This requires robust estimates of temporal process variance, for example in survival, and identification of weather covariates linked to interannual variability. 2. The European shag Phalacrocorax aristotelis (L.) shows unusually large variability in population size, and large-scale mortality events have been linked to winter gales. We estimated first-year, second-year and adult survival based on 43 years of ringing and dead recovery data from the Isle of May, Scotland, using recent methods to quantify temporal process variance and identify aspects of winter weather linked to survival. 3. Survival was highly variable for all age groups, and for second-year and adult birds process variance declined strongly when the most extreme year was excluded. Survival in these age groups was low in winters with strong onshore winds and high rainfall. Variation in first-year survival was not related to winter weather, and process variance, although high, was less affected by extreme years. A stochastic population model showed that increasing process variance in survival would lead to reduced population growth rate and increasing probability of extinction. 4. As in other cormorants, shag plumage is only partially waterproof, presumably an adaptation to highly efficient underwater foraging. We speculate that this adaptation may make individuals vulnerable to rough winter weather, leading to boom-and-bust dynamics, where rapid population growth under favourable conditions allows recovery from periodic large-scale weather-related mortality. 5. Given that extreme weather events are predicted to become more frequent, species such as shags that are vulnerable to such events are likely to exhibit stronger reductions in population growth than would be expected from changes in mean climate. Vulnerability to extreme events thus needs to be accounted for when predicting the ecological impacts of climate change.     

Some effects of inclement weather conditions on the survival and condition of bull-headed shrike nestlings

Inclement weather struck Japan in 1993, permitting a natural experiment for examining the effect of weather conditions on nesting success and nestling growth of the bull-headed shrike, Lanius bucephalus. Aspects of the breeding biology of bull-headed shrikes in relation to weather conditions and timing of breeding in 1992 and 1993 were examined. The two breeding seasons were divided into two periods, early and late, in each year. While the probability of nest survival in nestling stages during both periods was almost equal, the probability of nest survival in the egg stage during the early period was significantly lower than that during the late period. In 1993, the probability of nestling survival during the late period was significantly lower than during the early period; the late period had larger fluctuations of precipitation and was colder than the same period in 1992. The number of disappeared nestlings positively correlated with mean precipitation per day. The greater part of the disappeared nestlings was the lightest in each brood. Late breeders fledged lighter young than the early breeders. Although shrikes adopted hatching asynchrony, the late breeders could not surmount the unpredictable inclement weather in 1993.     

Impact of weather variation on feeding behaviour, nestling growth and brood survival in Wrynecks Jynx torquilla

While habitat destruction is thought to be the most important reason for the decline of the Wryneck Jynx torquilla throughout Europe, it has been hypothesised that weather factors may also play a crucial role. We studied the impact of variations in ambient temperature and rainfall on feeding behaviour, nestling growth and daily brood survival probabilities in a population from the Swiss Alps. We recorded the frequency of food provisioning and measured nestlings’ growth on 6 days at 17 nests, and monitored the success of 181 broods from 2002 to 2006. The mean feeding frequency was 7.9 feedings per hour, and increased with temperature and with nestling age. Changes of body mass, tarsus length and of the 8th primary feather were not strongly influenced by temperature and rainfall. Body growth was negatively influenced by the initial size of the different body measures towards the end of the growing phase of the corresponding body measure, pointing towards compensatory growth. Daily brood survival probabilities declined in the course of the season and with increasing rainfall, and they increased with increasing temperature and with increasing nestling age. Our findings highlight that the effects of adverse weather on feeding behaviour, nestling growth and nest survival in Wrynecks are complex, as they alter with the age of the nestlings and possibly with the duration of the adverse weather events. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorised users.     

Individual quality, survival variation and patterns of phenotypic selection on body condition and timing of nesting in birds

Questions about individual variation in quality and fitness are of great interest to evolutionary and population ecologists. Such variation can be investigated using either a random effects approach or an approach that relies on identifying observable traits that are themselves correlated with fitness components. We used the latter approach with data from 1,925 individual females of three species of ducks (tufted duck, Aythya fuligula; common pochard, Aythya ferina; northern shoveler, Anas clypeata) sampled on their breeding grounds at Engure Marsh, Latvia, for over 15 years. Based on associations with reproductive output, we selected two traits, one morphological (relative body condition) and one behavioral (relative time of nesting), that can be used to characterize individual females over their lifetimes. We then asked whether these traits were related to annual survival probabilities of nesting females. We hypothesized quadratic, rather than monotonic, relationships based loosely on ideas about the likely action of stabilizing selection on these two traits. Parameters of these relationships were estimated directly using ultrastructural models embedded within capture-recapture-band-recovery models. Results provided evidence that both traits were related to survival in the hypothesized manner. For all three species, females that tended to nest earlier than the norm exhibited the highest survival rates, but very early nesters experienced reduced survival and late nesters showed even lower survival. For shovelers, females in average body condition showed the highest survival, with lower survival rates exhibited by both heavy and light birds. For common pochard and tufted duck, the highest survival rates were associated with birds of slightly above-average condition, with somewhat lower survival for very heavy birds and much lower survival for birds in relatively poor condition. Based on results from this study and previous work on reproduction, we conclude that nest initiation date and body condition covary with both reproductive and survival components of fitness. These associations lead to a positive covariance of these two fitness components within individuals and to the conclusion that these two traits are indeed correlates of individual quality.     

The effects of survival and weather on lifetime egg production in a model damselfly

Abstract. 1. A simulation model is presented which describes, from field-estimated parameters, the effects of daily survival rate and proportion of sunny days on the lifetime egg production of females of the damselfly Coenagrion puella (L).
2. Lifetime egg production increases with daily survival rate and proportion of sunny days.
3. Estimates of mean lifetime egg production per female in bad and good summers in northern England ranged from 333 to 740.
4. The distribution, as well as the proportion, of sunny days influences lifetime egg production. For a given combination of daily survival and proportion of sunny days, lifetime egg production decreases as the distribution of sunny days becomes more clumped.
5. Lifetime egg production is largely determined by chance; females who begin their mature adult life during a period of sunny weather can produce many times more eggs than those whose mature adult life coincides with cloudy days.     

Monitoring of butterflies in semi-natural grasslands: diurnal variation and weather effects

The butterfly fauna was monitored in six semi-natural grasslands in southeastern Sweden. The aim was to evaluate monitoring criteria for wind, sunshine, temperature and time of day for butterfly species richness and abundances when using the line transect method. A total of 30,111 butterflies belonging to 46 species were recorded. Data from this study suggests somewhat stricter criteria for temperature and sunshine than stated in the widely used “Pollards walk”. A sharp decline in butterfly numbers were detected at temperatures below 19°C if the proportion of sunshine of the transect walk was below 80–85%. No effect of wind speed, up to five on the Beaufort scale, on observed numbers of species or individuals were found. Several butterfly species showed well-defined diurnal rythms of flight activity, and the results indicated that transect walks can be performed between −4.5 and +4 h from the time when the sun reached its highest point. The results of this study can be used to adjust general criteria in national monitoring and also for detailed regional and local monitoring where it may be important to adjust for diurnal rhythm and weather related bias.     

Additive and non-additive effects of simulated leaf and inflorescence damage on survival, growth and reproduction of the perennial herb Arabidopsis lyrata

Herbivores may damage both leaves and reproductive structures, and although such combined damage may affect plant fitness non-additively, this has received little attention. We conducted a 2-year field experiment with a factorial design to examine the effects of simulated leaf (0, 12.5, 25, or 50% of leaf area removed) and inflorescence damage (0 vs. 50% of inflorescences removed) on survival, growth and reproduction in the perennial herb Arabidopsis lyrata. Leaf and inflorescence damage negatively and independently reduced flower, fruit and seed production in the year of damage; leaf damage also reduced rosette size by the end of the first season and flower production in the second year. Leaf damage alone reduced the proportion of flowers forming a fruit and fruit production per plant the second year, but when combined with inflorescence damage no such effect was observed (significant leaf × inflorescence damage interaction). Damage to leaves (sources) caused a greater reduction in future reproduction than did simultaneous damage to leaves and inflorescences (sinks). This demonstrates that a full understanding of the effects of herbivore damage on plant fitness requires that consequences of damage to vegetative and reproductive structures are evaluated over more than 1 year and that non-additive effects are considered.     

Over-winter survival in subadult European rabbits: weather effects,density dependence,and the impact of individual characteristics

The survival probability of an individual may be limited by density-dependent mechanisms and by environmental stochasticity, but can also be modified by individual characteristics. In our study, we investigated over-winter survival of subadults of an enclosed European rabbit Oryctolagus cuniculus population in a temperate zone habitat over the period 1992–2002. We: (1) selected for appropriate models to explain individual variation in over-winter survival and the animals autumn body mass, the latter was used as a measure of the individual pre-winter body condition; and (2) aimed to compare the sensitivity of the target variables on the realised variation of the factors considered. Model selection based on information theory revealed that individual over-winter survival was best explained by the combination of autumn body mass, winter temperature, population density and sex, where the probability of survival was higher in females than in males. According to this model, the probability of survival reacted most sensitively to variation in the autumn body mass and in winter temperature. Individual autumn body mass was best explained by the combination of the date of birth, population density, and weather conditions by means of the percentage of rainy days during the first 2 months after the animals had emerged above ground, where the autumn body mass was negatively related to the percentage of rainy days. The chosen model suggested that the autumn body mass reacted most sensitively to variation in the date of birth. Combining these models, we found that weather conditions during two different periods of time as well as population density, sex and the date of birth operated together to determine the probability of over-winter survival. In particular, the study points out the high impact of environmental stochasticity on over-winter survival: (1) by direct effects of winter temperature conditions, and (2) by the indirect action of weather conditions to which the animals were exposed during the early period of juvenile development.     

Individual and spatio-temporal variations in the home range behaviour of a long-lived, territorial species

Despite the fact that investigations of home range behaviour have exponentially evolved on theoretical, analytical and technological grounds, the factors that shape animal home range behaviour still represent an unsolved question and a challenging field of research. However, home range studies have recently begun to be approached under a new integrated conceptual framework, considering home range behaviour as the result of the simultaneous influences of temporal, spatial and individual-level processes, with potential consequences at the population level. Following an integrated approach, we studied the influence of both external and internal factors on variations in the home range behaviour of 34 radiotagged eagle owl (Bubo bubo) breeders. Home range behaviour was characterised through complementary analysis of space use, movement patterns and rhythms of activity at multiple spatio-temporal scales. The effects of the different phases of the biological cycle became considerably evident at the level of movement patterns, with males travelling longer distances than females during incubation and nestling periods. Both external (i.e. habitat structure and composition) and internal (i.e. sex and health state) factors explained a substantial amount of the variation in home range behaviour. At the broader temporal scale, home range and core area size were negatively correlated with landscape heterogeneity. Males showed (1) smaller home range and core area sizes, (2) more complex home range internal structure and (3) higher rates of movement. The better the physiological condition of the individuals, the simpler the internal home range structure. Finally, inter- and intra-individual effects contributed to shaping space use and movement patterns during the biological cycle. Because of the plurality of behavioural and ecological processes simultaneously involved in home range behaviour, we claim that an integrative approach is required for adequate investigation of its temporal and spatial variation.     

Influence of drought and flowering on growth and water relations of perennial ryegrass populations

Spring-sown (vegetative) and autumn-sown (flowering) swards of Lolium perenne cvs Melle, Aurora and their hybrid growing in 0.9 m deep bins of soil in the glasshouse were compared, a) as drought progressed from June to August 1986, and b) after cutting, fertilising and re-watering.
During drought, vegetative plants produced more herbage, had initially higher leaf extension rates, had longer and wider leaves, maintained more tillers, had lower mortality, adjusted osmotic potential more effectively and had stomata less sensitive to stress than did flowering plants. On re-watering, previously stressed flowering plants regrew more slowly than the irrigated controls, whereas previously stressed vegetative plants regrew more rapidly than the controls.
Aurora was the most drought-resistant population, mainly because of low tiller death rates. The hybrid tended to have the lowest leaf water potentials and conductances and grew poorly during drought. Possible underlying physiological mechanisms are discussed.