Heterozygosity–behavior and heterozygosity–fitness correlations in a salamander with limited dispersal

Inbreeding and genetic drift can decrease genetic heterozygosity, and this low heterozygosity can depress fitness, resulting in heterozygosity–fitness correlations (HFCs). HFCs are typically small in magnitude, a result often attributed to power of the analyses. Animal behaviors often affect fitness and are often heritable to some degree. We hypothesized that heterozygosity influences behavior, which, in turn, potentially influences fitness. Specifically, in red-backed salamanders, Plethodon cinereus, which have limited dispersal and the potential for inbreeding, we tested whether heterozygosity, as estimated from six microsatellite loci, affected home range size, juvenile growth, and survival. We found that salamanders with higher heterozygosity had larger home ranges and grew faster, which is indicative of reproductive success. However, we found no effects of heterozygosity on survival. We conclude that, because activity in P. cinereus is tightly linked to food uptake and mass gain, heterozygosity influences growth via effects on foraging behavior. Future research should investigate how the relationship between heterozygosity, behavior, and fitness may be affected or mediated by endocrine or immune systems.     

How should we define fitness in structured metapopulation models? Including an application to the calculation of evolutionarily stable dispersal strategies

We define a fitness concept applicable to structured metapopulations consisting of infinitely many equally coupled patches. In addition, we introduce a more easily calculated quantity Rm that relates to fitness in the same manner as R0 relates to fitness in ordinary population dynamics: the Rm of a mutant is only defined when the resident population dynamics converges to a point equilibrium and Rm is larger (smaller) than 1 if and only if mutant fitness is positive (negative). Rm corresponds to the average number of newborn dispersers resulting from the (on average less than one) local colony founded by a newborn disperser. Efficient algorithms for calculating its numerical value are provided. As an example of the usefulness of these concepts we calculate the evolutionarily stable conditional dispersal strategy for individuals that can account for the local population density in their dispersal decisions. Below a threshold density x, at which staying and leaving are equality profitable, everybody should stay and above x everybody should leave, where profitability is measured as the mean number of dispersers produced through lines of descent consisting of non-dispersers.     

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     

Birthplace‐dependent dispersal: are directions of natal dispersal determined a priori?

Sib–sib or, more generally, family resemblance for dispersal seems a widespread characteristic of vertebrates, and the birthplace has the potential to shape the dynamics and features of animal populations. Dispersal studies have often stressed the fundamental link between the fate of dispersers and population dynamics, but few have focused on the dispersal directions of individuals, despite the profound implications that this may have on population distribution, structure, dynamics and viability. We investigated the directions followed by 72 radio‐tagged dispersers (43 males and 29 females from 14 nest sites) in an eagle owl Bubo bubo population, and assessed their a) inter‐individual distances during dispersal and b) age at dispersal departure. For siblings, as well as potential‐siblings (i.e. individuals born in the same nest in different years), the birthplace influenced inter‐individual distances and dispersal directions, i.e. dispersers from the same nest moved to similar locations during the study; moreover, in each year, individuals from the same birthplace moved across the same areas in a short time period. Finally, siblings and potential‐siblings born in the same nest in different years started dispersal at similar ages. Based on the movement patterns of dispersers we discuss: a) the potential implications of the birthplace‐dependent dispersal on the ideal free distribution theory, as well as in terms of kin competition, inbreeding avoidance and population dynamics; and, more generally, b) the effect of the temporal features of the natal dispersal on the concept of habitat suitability vs density of individuals developed by the ideal free distribution theory.     

Seed dispersal of East Asian coastal dune plants via seawater – short and long distance dispersal

Coastal plants provide precious and irreplaceable services to human and coastal ecosystems, but people are still rather unaware how coastal plants relate with sea tides. We assumed that their seeds may reflect some relationships with tides. Our objective was to understand seed dispersal of plants living in the upper coastal regions, not reached by normal tides, but inundated by storm surges. For this purpose, we observed seed characteristics, especially seed buoyancy, of coastal and non-inundated plants of the Shandong Peninsula, north China. Through field simulation, we studied how buoyancy affected the dispersal during tides, using twig segments as retrievable dummies of seeds, in order to collect evidences with concern, how seed dispersal could occur during storm surges, which are extremes of tides. Coastal plants had predominantly buoyant seeds, while inland-growing plants had largely non-buoyant seeds. Ninety percent of buoyant twig segments (mimicking seeds) were recaptured after tides on experimentally used coastal areas; only 1% of non-buoyant ones of such dummies were recaptured, whereas the rest probably had been swept away by wave action to deposit at the sea bottom. Buoyant twig segments (mimicking seeds) dispersed along the coastlines as tides surged towards the coast. About 10% of these buoyant twigs were swept away with off-shore currents, but they might land elsewhere, similar as it might happen with long-distance dispersal of seeds.     

Are mismatches the norm? Timing of flowering,fruiting, dispersal and germination and their fitness effects in Frangula alnus (Rhamnaceae)

The close morphological and temporal links between phases of plant growth and reproduction call for integrated studies incorporating several reproductive phases from flowering to recruitment, and associated plant‐animal interactions. Phenological strategies, as well as plastic phenological response to climate change, incorporate complex interactions between developmental constraints, pollination and seed dispersal. Relationships between reproductive phenology and components of fitness were studied for two years in the north‐temperate, self‐incompatible, insect‐pollinated, and bird‐dispersed shrub Frangula alnus (Rhamnaceae). Fruit set, dispersal, germination and juvenile survival, as well as seed mass and juvenile size were measured in relation to flowering, fruiting and germination time. The results suggest that effects of flowering and fruiting time prevailed in subsequent phases, to some extent as far as to the juvenile phase, but effects of timing were complex and had partly opposing effects on different fitness components. Early flowers had higher fruit‐set and experiments indicated that synchronous peak flowering increased fruit‐set, but later flowers had higher seed mass. Peak fruiting was not associated with peak dispersal. Late fruits derived from late flowers promoted dispersal. Juvenile recruitment was enhanced by increasing seed size. We conclude that the phenology of flowering and fruiting in F. alnus comprises several features, each with different and sometimes counteracting effects on fitness components. From a general perspective, this result implies that we should not expect to find finely tuned matches in timing specifically between flowering and pollinators, and fruiting and seed dispersing birds.     

Coevolution of dispersal in a parasitoid–host system

Interspecific interactions and the evolution of dispersal are both of interest when considering the potential impact of habitat fragmentation on community ecology, but the interaction between these processes is not well studied. We address this by considering the coevolution of dispersal strategies in a host–parasitoid system. An individual-based host–parasitoid metapopulation model was constructed for a patchy environment, allowing for evolution in dispersal rates of both species. Highly rarefied environments with few suitable patches selected against dispersal in both species, as did relatively static environments. Provided that parasitoids persist, all the variables studied led to stable equilibria in dispersal rates for both species. There was a tendency toward higher dispersal rates in parasitoids because of the asymmetric relationships of the two species to the patches: vacant patches are most valuable for hosts, but unsuitable for parasitoids, which require an established host population to reproduce. High host dispersal rate was favoured by high host population growth rate, and in the parasitoid by high growth rates in both species. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

When does the variance of replicator fitness decrease?

For the general replicator dynamics with regular relative advantage functions (having non-singular Jacobian and symmetrized Jacobian), it will be shown that the variance of marginal fitness of the replicators strictly decreases along each trajectory of the replicator dynamics near an interior rest point if and only if this rest point is a regular evolutionarily stable state.     

Geographic distributions of homosporous ferns: does dispersal obscure evidence of vicariance?

The central problem in biogeography is that interactions between different processes result in the formation of historical patterns, such that it is difficult to discriminate the relative roles of vicariance and dispersal. Ferns are distributed by small wind-dispersed propagules that are produced in very large numbers and capable of dispersing thousands of kilometers. Thus, most taxon distributions in ferns are assumed to be a function of dispersal rather than vicariance. Here, we review some case examples that provide good evidence for vicariance and dispersal in ferns. We then ask whether dispersal is so extensive in ferns that vicariance is no longer detectable in most cases. Although we think that too few studies have been carried out to make generalizations at this stage, we outline the criteria for an effective research programme that can address this issue. Phylogenetic and distributional data are needed, not only because they are lacking in an evolutionarily important group of organisms, but also because data from ferns and other cryptogams are likely to be crucial in making broad biogeographic statements.     

When is dispersal for dispersal? Unifying marine and terrestrial perspectives

Recent syntheses on the evolutionary causes of dispersal have focused on dispersal as a direct adaptation, but many traits that influence dispersal have other functions, raising the question: when is dispersal ‘for’ dispersal? We review and critically evaluate the ecological causes of selection on traits that give rise to dispersal in marine and terrestrial organisms. In the sea, passive dispersal is relatively easy and specific morphological, behavioural, and physiological adaptations for dispersal are rare. Instead, there may often be selection to limit dispersal. On land, dispersal is relatively difficult without specific adaptations, which are relatively common. Although selection for dispersal is expected in both systems and traits leading to dispersal are often linked to fitness, systems may differ in the extent to which dispersal in nature arises from direct selection for dispersal or as a by‐product of selection on traits with other functions. Our analysis highlights incompleteness of theories that assume a simple and direct relationship between dispersal and fitness, not just insofar as they ignore a vast array of taxa in the marine realm, but also because they may be missing critically important effects of traits influencing dispersal in all realms.     

Is farther seed dispersal better? Spatial patterns of offspring mortality in three rainforest tree species with different dispersal abilities

The paradigm that tropical trees with farther seed dispersal experience lower offspring mortality is currently based on within-species studies documenting higher survival of offspring located farther from conspecific adults and/or closer to light gaps. We determined whether the paradigm also holds among species by comparing spatial patterns of offspring mortality among three sympatric Neotropical rainforest tree species with the same seed dispersers but with different dispersal abilities. First, we assessed spatially non-random mortality for each species by measuring spatial shifts of the population recruitment curve (PRC) with respect to conspecific adults and light gaps across three early life stages: dispersed seeds, young seedlings and old seedlings. Then, we determined whether PRC shifts were greater for the species with short dispersal distances than for the species with greater dispersal distances. We found that the PRC shifted away from conspecific adults consistently across life stages, but we found no consistent PRC shifts towards gaps, suggesting that mortality was related more to the proximity of conspecifics than to absence of light gaps. PRC shifts away from adults were greatest in the species with the lowest dispersal ability, supporting the paradigm. Differential PRC shifts caused the spatial distribution of offspring to become almost independent with respect to adult trees and gaps in all three species, despite large differences in seed dispersal distance among these species. Our results provide direct empirical support for the paradigm that among tropical trees, species with farther seed dispersal are less impacted by spatially non-random mortality than are similar species with shorter dispersal distances. Thus, greater dispersal effectiveness merits extra investments of trees in seed dispersal ability, even at the cost of fecundity.     

Are red imported fire ants facilitators of native seed dispersal?

Invasive ants threaten native communities, in part, through their potential to disrupt mutualisms, yet invasive species may also facilitate native species. The red imported fire ant (Solenopsis invicta) is one of the most conspicuous invasive ants in North America and its high densities, combined with its potential to displace native ants, have led to concerns that it may disrupt ant-plant seed dispersal mutualisms. We examined the potential of fire ants to disperse seeds in the longleaf pine ecosystem by comparing the removal of elaiosome-bearing seeds by fire ants versus native ants. A total of 14 ant species were observed removing seeds, with fire ants responsible for more than half of all removals. While fire ants were the dominant seed remover in this system, they did not remove significantly more seeds than would be expected based on their population density (46% of ground-dwelling ants). Moreover, red imported fire ants were similar to native ants with respect to distance of seed movement and frequency of moving seeds back to the nest. Areas of higher fire ant densities were found to have greater rates of seed removal by ants without a subsequent drop in seed dispersal by native ants, suggesting that fire ant-invaded areas may experience overall higher levels of seed dispersal. Thus, fire ants may actually facilitate dispersal of elaiosome-bearing plant species in the longleaf pine ecosystem.     

How much is a lot? Seed dispersal by white-faced capuchins and implications for disperser-based studies of seed dispersal systems

The quantity of seeds dispersed is considered one of several means to determine the dispersal effectiveness of an animal. However, there is little consistency in the manner in which quantities are measured or presented. Here, we quantify seed dispersal by white-faced capuchin monkeys (Cebus capucinus) in Santa Rosa National Park, Costa Rica by measuring: degree of frugivory, number of plant species consumed, the number of seeds consumed per unit time, the number of seeds dispersed intact per unit time, and the number of seeds dispersed intact per unit space. Forty-nine percent of C. capucinus diet is composed of the fruit of 39 species, 4 of which constitute 82% of the frugivory. Seventy-four percent of consumed fruits contain seeds that pass intact through the capuchin digestive system. Capuchins pass a mean of 15.7 seeds of a mean of 1.3 species per defecation, and defecate 25.4 times per 12-h day. These numbers are compared with extant data for C. capucinus, and possible reasons for discrepancies among results between studies are discussed. We propose a standardization of quantitative measures of seed dispersal so that quantifications of seed dispersal can be compared within species, and eventually across species.     

Is cardiorespiratory fitness related to quality of life in survivors of breast cancer?

The purpose of this study was to investigate whether indices of cardiorespiratory fitness are related to quality of life (QOL) in women survivors of breast cancer. Using the European Organization for Research and Treatment of Cancer QLQ-30 questionnaire, we assessed the QOL of 16 participants (age, 50 +/- 9 years; body mass, 66.6 +/- 9.6 kg). All participants performed incremental cycle ergometer exercise to determine several indices of cardiorespiratory fitness (e.g., peak oxygen uptake [.V(O2)peak, in L.min(-1), ml.kg(-1).min(-1)]), peak power output (PPO, in W), PPO/ body mass (W.kg(-1), peak heart rate (HRpeak, b.min(-1), peak ventilation (VEpeak), and .V(O2) and heart rate (HR) at the ventilatory (VT) and respiratory compensation (RCT) thresholds. Relationships between QOL and variables were assessed using Spearman rank-difference correlation tests. A significant inverse relationship (p < 0.05) was found for QOL scores and values for age (years) and body mass (kg) ( = -0.53), %HRpeak@VT ( = -0.59) and %VEpeak@VT ( = -0.61). A significant positive relationship (p < 0.05) was found for QOL and PPO/body mass ( = 0.59) and HRpeak ( = 0.78), .V(O2)@RCT (ml.kg(-1.min(-1) ( = 0.51), power output (PO, expressed as either W or W.kg(-1) at RCT, and HR at RCT ( = 0.54). No other significant relationship was found between QOL and variables obtained from the tests. In conclusion, these findings highlight possible relationships between cardiorespiratory fitness and well-being in survivors of breast cancer. From a practical point of view, our data emphasize the need for this population to engage in programmed cardiorespiratory exercise training, mainly designed to improve VT and RCT. The improvement of both submaximal indices can have a beneficial effect on QOL.     

A ‘Goldilocks’ hypothesis for dispersal of biological control agents

The rate at which biological control agents disperse from release sites has important implications for their establishment and spread. Low rates of dispersal can yield spread that is too slow and may necessitate redistribution efforts for importation biological control and a high density of release sites for augmentation. Low dispersal rates may also lead to inbreeding at the site of release. On the other hand, high rates of dispersal can lead to Allee effects at the leading edge of the invasion front, potentially reducing the likelihood of establishment. Given these disadvantages associated with both low and high dispersal rates, we argue that intermediate rates of dispersal are likely to maximize the probability of establishment and appropriate spread for biological control agents released in the context of either importation or augmentative biological control. We consider this putative relationship a ‘Goldilocks hypothesis’ since it posits an optimum at intermediate values. In this review paper we begin by discussing the rationale for the Goldilocks hypothesis and then provide a case study from our work on importation biological control of the soybean aphid, Aphis glycines. Work on the soybean aphid parasitoid Binodoxys communis has shown that long-distance dispersal of immature parasitoids within winged migrating aphids is unlikely. This is likely good news for importation biological control because parasitoids dispersed in this manner would likely encounter crippling Allee effects. On the other hand, results from a field release study also suggest that female B. communis females (but not males) disperse actively from release sites. This female-biased dispersal may lead to strong mate-finding Allee effects and therefore may make establishment less likely.     

The heritability of fitness: Bad news for 'good genes'?

Some models of sexual selection depend on a female preference for 'good genes': females choose conspicuous males as these are advertising their possession of genes for fitness characters which can be inherited by their offspring. In contrast, Fisher's fundamental theorem of natural selection - which underlies much of population genetics theory - predicts that in a population at equilibrium there can be no additive genetic variation in fitness. Recent work on collared flycatchers in the wild shows that characters influencing fitness do indeed have a relatively low heritability. However, other studies of the inheritance of fitness in the laboratory suggest that under some circumstances a population may retain considerable genetic diversity for fitness characters. Genetically based female choice might hence have the potential to control the evolution of male sexual ornaments. More work on natural populations is needed; and birds may be a good place to start looking.     

A practical approximation of pollen dispersal–deposition for calculation of pollen productivity and quantification of vegetation

We compared simple mathematical pollen dispersal–deposition models with Gaussian plume models. The simple mathematical models proved equal or better approximations of real world pollen dispersal–deposition. We concluded that for most standard applications, such as estimating pollen productivity or quantitative vegetation reconstructions, simple mathematical models would perform satisfactory. Such easy-to-use models may lower the threshold to employ quantitative tools to palaeoecological questions.     

Do plant-community-based grazing regimes lead to epizoochorous dispersal of high proportions of target species?

The contribution of large-herbivore epizoochory to the transfer of seeds within and between areas is thought to be significant. But often seeds of ubiquitous species are dispersed, which may enhance ruderalization processes. In order to study the dispersal of target species by sheep, we employed a community-based grazing approach followed by intra- and inter-area sheep transfers (maximum transfer distance 3 km). In case of inter-area transfers, well-developed target communities of an open inland sand ecosystem are used as “source”, linked to less-developed sand habitats (“sink”) via sheep. Also other factors determining which species become dispersed under field conditions were tested: seed surface structure, seed mass, plant seed-releasing height and animal behaviour. Finally the influence of animal movement on seed detachment and the actual arrival of seeds within a “sink” were studied.

Sheep transfers resulted in the dispersal of 56 seed species, dominated by Red List (seven species) and other target species. Quantitatively, most transported seeds belonged to target species, whereas graminoid competitors were highly under-represented. Morphological traits enhance the attachment probability regardless of seed mass. But for seeds without these epizoochory-facilitating traits, mass seems to affect attachment negatively. Plant height affected the number of species present in sheep coats but not the seed quantities. Probably certain species in the vegetation produced large numbers of seeds, e.g. low-growing Medicago minima with seed surface structures and high-growing species Verbascum phlomoides without seed surface structures. Also, although transfer half-times were three times lower than grazing half-times, naturally attached Stipa capillata and Agrimonia procera seeds showed no significant detachment rates during transfer, whereas considerable losses were found during grazing. Other 3-km sheep transfers did not result in significant losses of either epizoochorously transported seed or species quantities. Our study shows that community-based grazing can lead to the dispersal of especially target species. Besides that, different habitat fragments can be connected to each other via sheep without significant seed losses along the way.     

Enhanced seed dispersal of Prunus africana in fragmented and disturbed forests?

Forest destruction and disturbance can have long-term consequences for species diversity and ecosystem processes such as seed dispersal. Understanding these consequences is a crucial component of conserving vulnerable ecosystems. In the heavily fragmented and disturbed Kakamega Forest, western Kenya, we studied seed dispersal of Prunus africana (Rosaceae). In the main forest, five forest fragments, and differently disturbed sites, we quantified the overall frugivore community as an indicator for species diversity. Furthermore, we determined the frugivores on 28 fruiting P. africana trees, estimated seed dispersal, crop size and the general fruit availability of surrounding trees. During the overall frugivore census we recorded 49 frugivorous species; 36 of them were observed visiting P. africana trees and feeding on their fruits. Although overall frugivore species richness was 1.1 times lower in fragments than in main forest sites and 1.02 times higher in highly disturbed than in less disturbed sites, P. africana experienced 1.1 times higher numbers of frugivores in fragments than in main forest sites and 1.5 times higher numbers of frugivores in highly disturbed than in less disturbed sites. Correspondingly, seed dispersal was 1.5 times higher in fragments than in main forest sites and 1.5 times higher in more disturbed than less disturbed sites. Fruit availability of surrounding trees and crop size influenced the number of visitors to some degree. Thus, the number of dispersed seeds seemed to be slightly higher in fragmented and highly disturbed sites. This indicates that loss of single species does not necessarily lead to a decrease of ecosystem services. However, loss of diversity could be a problem in the long term, as a multitude of species might act as buffer against future environmental change.