Fecundity limits in Frangula alnus (Rhamnaceae) relict populations at the species’ southern range margin

The geographic range of many temperate plant species is constrained by climate, but it remains little known how climate affects population performance at low-latitude range margins. This study investigated the reproduction of the Eurasian tree Frangula alnus in relict populations near its southwestern range limit in southern Spain. The aim was to identify the principal stages and causes of ovule loss experienced by these marginal populations. More than 6,800 flowers were monitored over 2 years, insect observations and different experiments were carried out to assess levels of pollen and resource limitation, as well as the influence of flowering phenology on seed production. Most ovule losses occurred during flower anthesis and were due to strong cross-pollen limitation. Fruit set was affected by tree size, light regime and flowering phenology, probably through their effects on pollinator behaviour. Fruit set was almost zero throughout the first half of the flowering season. Then it increased paralleling changes in pollinator abundance but was soon overridden by increasing ovule desiccation due to summer drought. Successful seed production was mostly confined to a brief period near the end of the flowering season. Adverse weather during this period in the second study year resulted in a threefold bud and flower mortality and a 50% decrease of fruit production. Spring rainfalls in southwestern Spain have diminished more than 30% through past decades leading to an earlier onset of summer drought. This trend and its adverse effects on seed production may contribute to explaining the recent decline of F. alnus at its southwestern range limit.Electronic Supplementary Material Supplementary material is available in the online version of this article at     

Is patience a virtue? Succession, light, and the death of invasive glossy buckthorn (Frangula alnus)

Non-native, invasive glossy buckthorn (Frangula alnus P. Mill.) threatens North American forests by inhibiting tree regeneration. While glossy buckthorn commonly invades younger, developing forests, we hypothesized that this species is competitively excluded as secondary succession proceeds. Specifically, we tested the hypothesis that glossy buckthorn mortality in New Hampshire forests is associated with low levels of photosynthetically active radiation (PAR), low levels of nutrients in the soil, and greater abundance of shade tolerant tree species. Twenty-six living and 26 dead glossy buckthorn individuals were randomly selected on three, mid-successional study sites in southeastern New Hampshire, USA. Shrub ages, indicated by wood ring counts, showed that both living and dead shrubs were members of the same cohort, roughly 16–18 years of age. Living and dead shrubs were compared as to (1) tree basal area in the surrounding forest (estimated by plot sampling), (2) light intensity in the vicinity of the buckthorn crowns (measured by Sunfleck Ceptometer), (3) canopy openness and % total transmitted PAR (estimated by hemispheric photography), and (4) concentrations of available Ca, P, Mg, and K in the B horizon. Relative to live buckthorn shrubs, dead individuals were associated with greater basal area of shade tolerant species, decreased PAR in the vicinity of shrub crown, and decreased soil concentrations of Ca, P, and Mg. The results are consistent with the hypothesis that competition for light and perhaps nutrients limits the ability of glossy buckthorn to persist in late successional stands. If these relationships are causal, active removal of glossy buckthorn might not be required to reduce its abundance in late successional stands.     

Estimating fitness consequences of dispersal: a road to 'know-where'? Non-random dispersal and the underestimation of dispersers' fitness

1. Many studies investigating fitness correlates of dispersal in vertebrates report dispersers to have lower fitness than philopatric individuals. However, if dispersers are more likely to produce dispersing young or are more likely to disperse again in the next year(s) than philopatric individuals, there is a risk that fitness estimates based on local adult survival and local recruitment will be underestimated for dispersers. 2. We review the available empirical evidence on parent-offspring resemblance and individual lifelong consistency in dispersal behaviour, and relate these studies to recent studies of fitness correlates of dispersal in vertebrates. 3. Of the 12 studies testing directly for parent-offspring resemblance in dispersal propensity, five report a significant resemblance. The average effect size (r) of parent-offspring resemblance in dispersal was 0.15 [95% confidence interval (CI) = 0.07-0.22], with no difference between the sexes (average weighted effect size of 0.12 (0.08-0.16) and 0.16 (0.11-0.20) for females and males, respectively). Only three studies report data on within-individual consistency in dispersal propensity, of which two suggest dispersers to be more likely to disperse again. 4. To assess the magnitude of fitness underestimation expected for dispersing individuals depending on the heritability of dispersal distance and study area size, we used a simulation approach. Even when study area size is 10 times the mean dispersal distance, local recruitment per breeding event may be underestimated by 4-10%, generating a potential difference of 4-60% in average lifetime production of recruits between dispersing and philopatric individuals, with larger differences in long-lived species. 5. Estimates of both fitness correlates of dispersal and parent-offspring resemblance or within-individual consistency in dispersal behaviour have been reported for 11 species. Although some comparisons suggest genuine differences in fitness components between philopatric and dispersing individuals, others, based on adult and juvenile survival, are open to the alternative explanation of biased fitness estimates. 6. We list three potential ways of reducing the risk of making wrong inferences on biased fitness estimates due to such non-random dispersal behaviour between dispersing and philopatric individuals: (a) diagnosing effects of non-random dispersal, (b) reducing the effects of spatially limited study area and (c) performing controlled experiments.     

Modified dispersal‐related traits in disjunct populations of bird‐dispersed Frangula alnus (Rhamnaceae): a result of its Quaternary distribution shifts?

Many European tree species survived Pleistocene glaciations in Mediterranean refugia and rapidly recolonized temperate Europe afterwards. Inter‐ and postglacial migration processes are assumed to have catalized evolutionary optimizations of dispersal‐related traits, but up to now empirical evidence is lacking in vertebrate‐dispersed plants. We investigated if south Iberian glacial relict and central European “colonizer” populations of the bird‐dispersed tree Frangula alnus have experienced differentiations of dispersal‐related traits which increase the mobility of northern populations. A comparison of lifetime reproductive strategy, disperser guilds, ripening phenology, and fruit design revealed considerable differences. Compared to south Iberian conspecifics, central European plants were considerably smaller and experienced a highly accelerated generation turnover. In south Iberian populations seed dispersal was carried out almost completely by resident birds which occurred in constant abundances throughout the ripening season. In contrast, central European seeds were dispersed by migrants whose abundances changed considerably during the ripening season. Several bird species were involved in both study areas but rendered different importance for seed dispersal. The fruit ripening pattern was highly asynchronous throughout the ripening season in south Iberia, while central European trees showed a complex ripening sequence which resulted in a significant correlation between fruit abundance and changing disperser availability. Central European fruits were smaller and showed a considerably smaller seed load than south Iberian fruits, thus presumably being more attractive for their small‐sized main dispersers (Sylvia warblers). Chemical analyses revealed significant differences in contents of water, glucose, fructose, proteins, ash. and phenolic compounds. The extensive differentiation of dispersal‐related traits in F. alnus suggests that even weak selective pressures by frugivores may induce evolutionary adjustments of dispersal traits over large time scales. We suggest that the differences we observe today evolved during the species' distribution shifts in the Quaternary.     

Intraspecific variation in fruit–frugivore interactions: effects of fruiting neighborhood and consequences for seed dispersal

The extent of specialization/generalization continuum in fruit–frugivore interactions at the individual level remains poorly explored. Here, we investigated the interactions between the Neotropical treelet Miconia irwinii (Melastomataceae) and its avian seed dispersers in Brazilian campo rupestre. We built an individual-based network to derive plant degree of interaction specialization regarding disperser species. Then, we explored how intraspecific variation in interaction niche breadth relates to fruit availability on individual plants in varying densities of fruiting conspecific neighbors, and how these factors affect the quantity of viable seeds dispersed. We predicted broader interaction niche breadths for individuals with larger fruit crops in denser fruiting neighborhoods. The downscaled network included nine bird species and 15 plants, which varied nearly five-fold in their degree of interaction specialization. We found positive effects of crop size on visitation and fruit removal rates, but not on degree of interaction specialization. Conversely, we found that an increase in the density of conspecific fruiting neighbors both increased visitation rate and reduced plant degree of interaction specialization. We suggest that tracking fruit-rich patches by avian frugivore species is the main driver of density-dependent intraspecific variation in plants’ interaction niche breadth. Our study shed some light on the overlooked fitness consequences of intraspecific variation in interaction niches by showing that individuals along the specialization/generalization continuum may have their seed dispersed with similar effectiveness. Our study exemplifies how individual-based networks linking plants to frugivore species that differ in their seed dispersal effectiveness can advance our understanding of intraspecific variation in the outcomes of fruit–frugivore interactions.

     

Seed dispersal in Cakile edentula var. lacustris : decoupling the fitness effects of density and distance from the home site

A factorial design of three densities of siblings at three local distances from seed parents was employed to distinguish effects of density from effects of dispersal distance on lifespan and fruit production of Cakile edentula var. lacustris, a plant with heteromorphic seeds. The segmented fruits produce two seed types: proximal and distal, with distal seeds having greater mass and greater dispersibility. Effects of longer distances (0.5 km and 30 km) on lifespan and fruit production were investigated using plants at low density. The prediction was tested that the greater seed mass of distal seeds increases fitness when seeds are dispersed into sites of unknown quality away from the home site or when seeds are dispersed to low density. High density caused earlier mortality and lower probability of reproduction. Distance from the maternal plant did not influence lifespan or reproduction at distances of 15 m or less, but lifespan was longer 0.5 km from the home site. No interaction was detected between the effects of density and distance on either lifespan or total fitness. Environmental conditions that influence fitness did not vary as a function of dispersal distance in this system, and favorable conditions at the home site did not persist between generations. Therefore, selection on dispersion patterns in natural conditions is likely to be through effects of density rather than dispersal distance. Proximal seeds had greater reproduction than distal seeds at the home site, and distal seeds had greater reproduction at the more distant sites (but not the most distant site), as expected, but these performance differences could not be attributed to differences in mass between the two seed types. Reduced seed mass was favored at the most distant site, but larger seed mass was favored most strongly at low density. Seeds that are dispersed to low density are larger, suggesting that although kin selection may limit the effectiveness of individual selection to increase seed mass under conditions of sibling competition, density-dependent individual selection on seed mass, rather than distant-dependent selection, also contributes to the observed associations among seed type, seed mass and dispersal ability. Received: 21 October 1996 / Accepted: 4 December 1996     

“Perchance to dream?”: Assessing the effects of dispersal strategies on the fitness of expanding populations

Unraveling the patterns of animals’ movements is crucial to understanding the basics of biogeography, tracking range shifts resulting from climate change, and predicting and preventing biological invasions. Many researchers have modeled animals’ dispersal under the assumptions of various movement strategies, either predetermined or directed by external factors, but none have compared the effects of different movement strategies on population survival and fitness. In this paper, using an agent-based model with a landscape divided into cells of varying quality, we compare the ecological success of three movement and habitat selection strategies (MHSSs): (i) Smart, in which animals choose the locally optimal cell; (ii) Random, in which animals move randomly between cells without taking into account their quality; (iii) Dreamer, in which animals attempt to find a habitat of dream whose quality is much higher than that of the habitat available on the map. We compare the short-term success of these MHSSs in good, medium and bad environments. We also assess the effect of temporal variation of habitat quality (specifically, winter harshness) on the success of each MHSS. Success is measured in terms of survival rate, dispersal distance, accumulated energy and quality of settled habitat. The most general conclusion is that while survival rate, accumulated energy and quality of settled habitat are affected primarily by overall habitat composition (proportions of different habitat types in the landscape), dispersal distance depends mainly on the MHSS. In medium and good environments, the Dreamer strategy is highly successful: it simultaneously outperforms the Smart strategy in dispersal distance and the Random strategy in terms of the other metrics.     

Are mutations usually deleterious? A perspective on the fitness effects of mutation accumulation

Evolutionary Ecology - All adaptive alleles in existence today began as mutations, but a common view in ecology, evolution, and genetics is that non-neutral mutations are much more likely to be...     

Influence of primitive Biłgoraj horses on the glossy buckthorn (Frangula alnus)-dominated understory in a mixed coniferous forest

Changes in the understory dominated by glossy buckthorn Frangula alnus via the influence of primitive horses were analyzed in a 28-year-old enclosure in the village of Szklarnia at the Biłgoraj Horse-Breeding Centre near Janów Lubelski (eastern Poland). The analysis was conducted in 20 circular plots (30 m²) defined in adjacent, similar forest stands (enclosed and control). Disturbance by the horses, mainly through trampling, caused numerous paths to form within the glossy buckthorn-dominated understory and led to a decrease in density of stems of lower height classes (30–80 and 81–130 cm, respectively). An increase in species diversity at the expense of glossy buckthorn density was also observed. The horses' trampling caused an increase in Padus avium density and the encroachment of other woody plant species that were less shade-tolerant and grew well in soils rich in nutrients. An increase in the density of woody plants over 180 cm above ground was observed within the enclosure, which was probably the result of the horses’ excretion of feces. The results presented here provide new insight into the ecological role that horses play in forest-meadow landscape mosaics, which, via altering the development of vegetation, may contribute to an increase in biodiversity within forest habitats.     

Field evidence of dispersal of branchiopods, ostracods and bryozoans by teal (Anas?crecca) in the Camargue (southern France)

Invertebrate propagules may survive internal and external transports by waterbirds, thus facilitating their dispersal between aquatic habitats. However, field data on such transport remain limited, especially for exozoochory. We quantified and compared the rates of internal and external invertebrate transports simultaneously in a wintering population of teal (Anas crecca) in the Camargue (southern France). We inspected lower gut (rectum) contents of birds that had been shot (N = 366) and washed birds that had been live-trapped (N = 68) during the winters 2006–2007 and 2007–2008. At least one propagule was recorded in 2.5% and 10.3% of internal and external samples, respectively. Cladoceran, ostracod, plumatellid bryozoan and anostracan propagules were all recorded in both internal and external samples. Hatching confirmed their viability, except for anostracan eggs. At least three cladoceran species and two ostracod species were recorded, none of which had previously been shown to be dispersed by birds. Amongst external samples, we recorded a significant seasonal trend in 1 year with most propagules recorded before December, keeping with a seasonal decline in the availability of propagules on the water surface. This study provides evidence that teal may be important vectors of invertebrate dispersal both within the Camargue and along migratory flyways.     

Integration of flowering dates in phenology and pollen counts in aerobiology: analysis of their spatial and temporal coherence in Germany (1992–1999)

We studied the possibility of integrating flowering dates in phenology and pollen counts in aerobiology in Germany. Data were analyzed for three pollen types (Betula, Poaceae, Artemisia) at 51 stations with pollen traps, and corresponding phenological flowering dates for 400 adjacent stations (< 25 km) for the years 1992–1993 and 1997–1999. The spatial and temporal coherence of these data sets was investigated by comparing start and peak of the pollen season with local minima and means of plant flowering. Our study revealed that start of birch pollen season occurred on average 5.7 days earlier than local birch flowering. For mugwort and grass, the pollen season started on average after local flowering was observed; mugwort pollen was found 4.8 days later and grass pollen season started almost on the same day (0.6 days later) as local flowering. Whereas the peak of the birch pollen season coincided with the mean flowering dates (0.4 days later), the pollen peaks of the other two species took place much later. On average, the peak of mugwort pollen occurred 15.4 days later than mean local flowering, the peak of grass pollen catches followed 22.6 days after local flowering. The study revealed a great temporal divergence between pollen and flowering dates with an irregular spatial pattern across Germany. Not all pollen catches could be explained by local vegetation flowering. Possible reasons include long-distance transport, pollen contributions of other than phenologically observed species and methodological constraints. The results suggest that further research is needed before using flowering dates in phenology to extrapolate pollen counts.     

Are there latitudinal and altidudinal Rapoport effects in the geographic ranges of Andean passerine birds?

We analysed the range-sizes of 835 Andean passerine species (including 414 endemics and 421 non-endemics) to test for latitudinal and altitudinal Rapoport effects (LRE and ARE). We tested for positive range-size: latitude/altitude correlations using three different methods: (i) Rohde's mid-point method, (ii) species sorted out by altitude, and (iii) a phylogenetic comparative method (CAIC). Using Rohde's mid-point method, the mean latitudinal extent of species does not follow a Rapoport pattern, but the mean latitudinal occupancy of all passerines and non-endemics do increase with latitude. The latitudinal ranges of endemics sorted out by altitude follow a reverse Rapoport effect, but non-endemics support the pattern. CAIC confirms the latitudinal increase in the occupancy of non-endemics, but regressions have low coefficients of determination. The ARE is supported by the mean altitudinal extent of species, but the trend vanishes when controlling for geometric effects. Low-altitude species occupy about the same proportion of the available altitudinal space as do high-altitude ones. Our analyses suggest that latitude and altitude have low explanatory power for understanding the spatial variation in range-sizes at a continental scale. We show how different patterns can emerge from applying different criteria to the analysis of data.     

Evidence of opposing fitness effects of parental heterozygosity and relatedness in a critically endangered marine turtle?

How individual genetic variability relates to fitness is important in understanding evolution and the processes affecting populations of conservation concern. Heterozygosity–fitness correlations (HFCs) have been widely used to study this link in wild populations, where key parameters that affect both variability and fitness, such as inbreeding, can be difficult to measure. We used estimates of parental heterozygosity and genetic similarity (‘relatedness’) derived from 32 microsatellite markers to explore the relationship between genetic variability and fitness in a population of the critically endangered hawksbill turtle, Eretmochelys imbricata. We found no effect of maternal MLH (multilocus heterozygosity) on clutch size or egg success rate, and no single‐locus effects. However, we found effects of paternal MLH and parental relatedness on egg success rate that interacted in a way that may result in both positive and negative effects of genetic variability. Multicollinearity in these tests was within safe limits, and null simulations suggested that the effect was not an artefact of using paternal genotypes reconstructed from large samples of offspring. Our results could imply a tension between inbreeding and outbreeding depression in this system, which is biologically feasible in turtles: female‐biased natal philopatry may elevate inbreeding risk and local adaptation, and both processes may be disrupted by male‐biased dispersal. Although this conclusion should be treated with caution due to a lack of significant identity disequilibrium, our study shows the importance of considering both positive and negative effects when assessing how variation in genetic variability affects fitness in wild systems.     

In vitro and in vivo comparisons of the effects of the fruiting body and mycelium of Antrodia camphorata against amyloid β-protein-induced neurotoxicity and memory impairment

Antrodia camphorata is a particular and precious medicinal mushroom, and its fruiting body was found to provide more efficient protection from oxidative stress and inflammation than its mycelium because of its higher content of triterpenoids, total phenols, and so on. In the previous in vitro studies, the mycelium of A. camphorata is proven to provide strong neuroprotection in neuron cells and suggested to have the potential of protection against neurotoxicity of amyloid β-protein (Aβ) known as the risk factor toward Alzheimer's disease (AD) development. However, the in vivo study and the comparison study with the fruiting body have not yet been investigated. This study compared the effect of the fruiting body and mycelium of A. camphorata on alleviating the Aβ40-induced neurocytotoxicity in the in vitro Aβ-damaged neuron cell model (PC-12 cell treated with Aβ40) and memory impairment in the in vivo AD animal model induced with a continuous brain infusion of Aβ40. In the results of in vitro and in vivo studies, the fruiting body possessed stronger anti-oxidative and anti-inflammatory abilities for inhibiting neurocytotoxicity in Aβ40-treated PC-12 cells and Aβ40 accumulation in Aβ40-infused brain than mycelium. Moreover, hyperphosphorylated tau (p-tau) protein expression, known as an important AD risk factor, was suppressed by the treatment of fruiting body rather than that of mycelium in the in vitro and in vivo studies. These comparisons supported the reasons why the fruiting body resulted in a more significant improvement effect on working memory ability than mycelium in the AD rats.     

Do different parasite species interact in their effects on host fitness? A case study on parasites of the amphipod Paracalliope fluviatilis

There is a gap in our understanding of the relative and interactive effects of different parasite species on the same host population. Here we examine the effects of the acanthocephalan Acanthocephalus galaxii, an unidentified cyclophyllidean cestode, and the trematodes Coitocaecum parvum and Microphallus sp. on several fitness components of the amphipod Paracalliope fluviatilis, using a combination of infection surveys and both survival and behavioural trials. In addition to significant relationships between specific parasites and measures of amphipod survival, maturity, mating success and behaviour, interactions between parasite species with respect to amphipod photophilia were also significant. While infection by either A. galaxii or C. parvum was associated with increased photophilia, such increases were negated by co-infection with Microphallus sp. We hypothesize that this is due to the more subtle manipulative effect of A. galaxii and C. parvum being impaired by Microphallus sp. We conclude that the low frequency at which such double infections occur in our sampled population means that such interactions are unlikely to be important beyond the scale of the host individual. Whether or not this is generally true, implying that parasitological models and theory based on single parasite species studies do generally hold, requires cross-species meta-analytical studies.     

Variation in timing and abundance of flowering by Delphinium barbeyi Huth (Ranunculaceae): the roles of snowpack, frost, and La Niña, in the context of climate change

Delphinium barbeyi is a common herbaceous wildflower in montane meadows at 2,900 m near the Rocky Mountain Biological Laboratory, and its flowers are important nectar resources for bumblebees and hummingbirds. During the period 1977-1999 flowering was highly variable in both timing (date of first flower ranged from 5 July to 6 August, mean=17 July) and abundance (maximum open flowers per 2Ƕ-m plot ranged from 11.3 to 197.9, mean=82). Time and abundance of flowering are highly correlated with the previous winter's snowpack, as measured by the amount of snow remaining on the ground on 15 May (range 0-185 cm, mean=67.1). We used structural equation modeling to investigate relationships among snowpack, first date of bare ground, first date of flowering, number of inflorescences produced, and peak number of flowers, all of which are significantly correlated with each other. Snowpack depth on 15 May is a significant predictor of the first date of bare ground (R²=0.872), which in turn is a significant predictor of the first date of flowering (R²=0.858); snowpack depth is also significantly correlated with number of inflorescences produced (R²=0.713). Both the number of inflorescences and mean date of first flowering are significant predictors of flowers produced (but with no residual effect of snowpack). Part of the effect of snowpack on flowering may be mediated through an increased probability of frost damage in years with lower snowpack - the frequency of early-season "frost events" explained a significant proportion of the variance in the number of flowers per stem. There is significant reduction of flower production in La Niña episodes. The variation in number of flowers we have observed is likely to affect the pollination, mating system, and demography of the species. Through its effect on snowpack, frost events, and their interaction, climate change may influence all of these variables.     

Flowering and fruiting phenology in the coastal shrublands of Doñana,south Spain

Flowering and fruiting phenological patterns at the individual-, population-, and community level were studied in a southern Spanish scrub community composed of 30 shrub species. Few individuals of any species produced a high number of flowers. Intrapopulation deviation in the peak time of flowering showed a strong and positive skewness. Relative flowering duration, however, displayed a virtually normal distribution. Generally, species flowering in spring have a short flowering time, while species flowering earlier or later in the year show significantly longer flowering periods. Species were in bloom throughout the year, but there was a major peak during spring and two lesser ones in autumn and early summer. Shallow rooting taxa in typically mediterranean genera displayed a strategy of spring flowering and summer fruiting. Summer and autumn flowering occurred among heath-like shrubs of relatively wet places, and forest-associated, vertebrate-dispersed species which commonly have underground storage organs. Species with ripe fruits presented two peaks, the major one during the summer including the majority of taxa with seeds dispersed by non-vertebrate agents. There was a minor fruiting peak in autumn dominated by taxa that rely on vertebrates for dispersal. The complex seasonal patterns observed are interpreted in relation to environmental conditions and physiological constraints on species living in a highly seasonal climate.