Environmental variability and semelparity vs. iteroparity as life histories

Research on the evolution of life histories addresses the topic of fitness trade-offs between semelparity (reproducing once in a lifetime) and iteroparity (repeated reproductive bouts per lifetime). Bulmer (1994) derived the relationship v+P(A)<1 (P(A) is the adult survival;vb(S) and b(S) are the offspring numbers for iteroparous and semelparous breeding strategies, respectively), under which a resident semelparous population cannot be invaded by an iteroparous mutant when the underlying population dynamics are stable. We took Bulmer's population dynamics, and added noise in juvenile and adult survival and in offspring numbers. Long-term coexistence of the two strategies is possible in much of the parameter region ofv +P(A)<1 when noise occurs simultaneously in all three components, or (more restricted) when it affects juvenile and adult survival or adult survival and offspring numbers. Iteroparity cannot persist when the environmental variability involves juvenile survival and offspring numbers, or when the noise acts on the three components separately.     

Between semelparity and iteroparity: Empirical evidence for a continuum of modes of parity

The number of times an organism reproduces (i.e., its mode of parity) is a fundamental life‐history character, and evolutionary and ecological models that compare the relative fitnesses of different modes of parity are common in life‐history theory and theoretical biology. Despite the success of mathematical models designed to compare intrinsic rates of increase (i.e., density‐independent growth rates) between annual‐semelparous and perennial‐iteroparous reproductive schedules, there is widespread evidence that variation in reproductive allocation among semelparous and iteroparous organisms alike is continuous. This study reviews the ecological and molecular evidence for the continuity and plasticity of modes of parity—that is, the idea that annual‐semelparous and perennial‐iteroparous life histories are better understood as endpoints along a continuum of possible strategies. I conclude that parity should be understood as a continuum of different modes of parity, which differ by the degree to which they disperse or concentrate reproductive effort in time. I further argue that there are three main implications of this conclusion: (1) that seasonality should not be conflated with parity; (2) that mathematical models purporting to explain the general evolution of semelparous life histories from iteroparous ones (or vice versa) should not assume that organisms can only display either an annual‐semelparous life history or a perennial‐iteroparous one; and (3) that evolutionary ecologists should base explanations of how different life‐history strategies evolve on the physiological or molecular basis of traits underlying different modes of parity.     

Comparative phylogenetic analysis of the evolution of semelparity and life history in salmonid fishes

The selective pressures involved in the evolution of semelparity and its associated life-history traits are largely unknown. We used species-level analyses, independent contrasts, and reconstruction of ancestral states to study the evolution of body length, fecundity, egg weight, gonadosomatic index, and parity (semelparity vs. degree of iteroparity) in females of 12 species of salmonid fishes. According to both species-level analysis and independent contrasts analysis, body length was positively correlated with fecundity, egg weight, and gonadosomatic index, and semelparous species exhibited a significantly steeper slope for the regression of egg weight on body length than did iteroparous species. Percent repeat breeding (degree of iteroparity) was negatively correlated with gonadosomatic index using independent contrasts analysis. Semelparous species had significantly larger eggs by species-level analysis, and the egg weight contrast for the branch on which semelparity was inferred to have originated was significantly larger than the other egg weight contrasts, corresponding to a remarkable increase in egg weight. Reconstruction of ancestral states showed that egg weight and body length apparently increased with the origin of semelparity, but fecundity and gonadosomatic index remained more or less constant or decreased. Thus, the strong evolutionary linkages between body size, fecundity, and gonadosomatic index were broken during the transition from iteroparity to semelparity. These findings suggest that long-distance migrations, which increase adult mortality between breeding episodes, may have been necessary for the origin of semelparity in Pacific salmon, but that increased egg weight, leading to increased juvenile survivorship, was crucial in driving the transition. Our analyses support the life-history hypotheses that a lower degree of repeat breeding is linked to higher reproductive investment per breeding episode, and that semelparity evolves under a combination of relatively high juvenile survivorship and relatively low adult survivorship.     

Costs of reproduction can explain the correlated evolution of semelparity and egg size: theory and a test with salmon

Species’ life history traits, including maturation age, number of reproductive bouts, offspring size and number, reflect adaptations to diverse biotic and abiotic selection pressures. A striking example of divergent life histories is the evolution of either iteroparity (breeding multiple times) or semelparity (breed once and die). We analysed published data on salmonid fishes and found that semelparous species produce larger eggs, that egg size and number increase with salmonid body size among populations and species and that migratory behaviour and parity interact. We developed three hypotheses that might explain the patterns in our data and evaluated them in a stage‐structured modelling framework accounting for different growth and survival scenarios. Our models predict the observation of small eggs in iteroparous species when egg size is costly to maternal survival or egg number is constrained. By exploring trait co‐variation in salmonids, we generate new hypotheses for the evolution of trade‐offs among life history traits.     

How competitive intransitivity and niche overlap affect spatial coexistence

Competitive intransitivity is mostly considered outside the main body of coexistence theories that rely primarily on the role of niche overlap and differentiation. How the interplay of competitive intransitivity and niche overlap jointly affects species coexistence has received little attention. Here, we consider a rock–paper–scissors competition system where interactions between species can represent the full spectra of transitive–intransitive continuum and niche overlap/differentiation under different levels of competition asymmetry. By comparing results from pair approximation that only considers interference competition between neighbouring cells in spatial lattices, with those under the mean-field assumption, we show that 1) species coexistence under transitive competition is only possible at high niche differentiation; 2) in communities with partial or pure intransitive interactions, high levels of niche overlap are not necessary to beget species extinction; and 3) strong spatial clustering can widen the condition for intransitive loops to facilitate species coexistence. The two mechanisms, competitive intransitivity and niche differentiation, can support species persistence and coexistence, either separately or in combination. Finally, the contribution of intransitive loops to species coexistence can be enhanced by strong local spatial correlations, modulated and maximised by moderate competition asymmetry. Our study, therefore, provides a bridge to link intransitive competition to other generic ecological theories of species coexistence.     

Dispersal and spatial heterogeneity allow coexistence between enemies and protective mutualists

Protective mutualisms, where a symbiont reduces the negative effects of another species on a shared host, represent a common type of species interaction in natural communities, yet it is still unclear what ecological conditions might favor their emergence. Studies suggest that the initial evolution of protective mutualists might involve closely related pathogenic variants with similar life histories, but different competitive abilities and impacts on host fitness. We derive a model to evaluate this hypothesis and show that, in general, a protective variant cannot spread from rarity or exclude a more pathogenic strain. While the conditions allowing mutualist invasion are more likely with increased environmental productivity, they still depend on initial densities in the invaded patch exceeding a threshold, highlighting the likely importance of spatial structure and demographic stochasticity. Using a numerical simulation approach, we show that regional coexistence is in fact possible in an explicitly spatial system and that, under some circumstances, the mutualist population can exclude the enemy. More broadly, the establishment of protective mutualists may be favored when there are other life‐history differences from more pathogenic symbionts, such as vertical transmission or additional direct benefits to hosts.     

Integrating nested spatial scales: implications for the coexistence of competitors on a patchy resource

1. Intraspecific aggregation at a single spatial scale can promote the coexistence of competitors. This paper demonstrates how this same mechanism can be applied to the many systems that are patchy at two scales, with patches nested within 'superpatches'.
2. Data are presented from a field study showing that insects living in rotting fruits have aggregated distributions in the fruits under a single tree, and that the mean density and degree of aggregation varies significantly among trees. Observations in this system motivate the following models.
3. A model of competition has been developed between two species which explicitly represents spatial variation at two scales. By integrating the probability distributions for each scale, the marginal distributions of competitors over all patches can be found and used to calculate coexistence criteria. This model assumes global movement of the competitors.
4. Although spatial variation at a single scale may not be sufficient for coexistence, the total variation over all patches can allow coexistence. Variation in mean densities among superpatches and variation in the degree of aggregation among superpatches both promote coexistence, but act in different ways.
5. A second model of competition between two species is described which incorporates the effects of limited movement among superpatches. Limited movement among superpatches generally promotes coexistence, and also leads to correlations among aggregation and the mean densities of competitors.     

Pattern of gonad maturation and the question of semelparity in the paedomorphic goby Aphia minuta

At variance with previous reports, the paedomorphic goby Aphia minuta does not show semelparity, but abbreviate iteroparity like other small Mediterranean gobiid species. In spring, the mature ovary contains several batches of eggs at different stages of vitellogenesis. This indicates that the breeding season of A. minuta is quite long and that spawning takes place at least twice during its short lifespan, involving in each ovulatory wave different batches of oocytes which undergo vitellogenesis at different times. These observations agree with the testicular cycle data. In males, which in spring exhibit active spermiation coinciding with deposition of the first batch of oocytes, spermatogonial mitosis also resumes, eventually resulting in the production of new sperm in summer.     

Effects of Ration and Temperature on Reproduction of a Semelparous and an Iteroparous Triclad

Summary

This paper compares aspects of the reproduction of a semelparous triclad, Dendrocoelum lacteum, with those of an iteroparous triclad, Dugesia polychroa. Cocoon output was less sensitive to ration and possibly temperature for the former as compared with the latter species. Cocoons of D. lacteum were on average bigger and contained more hatchlings than those of D. polychroa. The sizes of hatchlings did not differ between species, but for D. lacteum only, the hatchlings emerging from late cocoons were significantly larger, and less numerous, than those emerging from earlier cocoons. Possible reasons for these differences are considered.     

The geometry of coexistence

Understanding the processes that maintain diversity has been the focus of extensive study, yet there is much that has not been integrated into a cohesive framework. First, there is a separation of perspective. Ecological and evolutionary approaches to diversity have progressed in largely parallel directions. Second, there is a separation of emphasis. In both ecology and population genetics, classical theories favour local explanations with emphasis on population dynamics and selection within populations, while contemporary theories favour spatial explanations, with emphasis on population structure and interactions among populations. What is lacking is a comparative approach that evaluates the relative importance of local and spatial processes in maintaining genetic and ecological diversity. I present a framework for diversity maintenance that emphasizes the comparative approach. I use a well-known but little-used mathematical approach, the perturbation theorem for dynamical systems, to identify key points of contact between ecological and population genetic theories of coexistence. These connections provide for a synthesis of several important concepts: population structure (source-sink versus extinction-colonization), spatial heterogeneity (intrinsic versus extrinsic) in fitness and competitive ability, and temporal scales over which local and spatial processes influence diversity. This framework ties together a large and diverse body of theory and data from ecology and population genetics. It yields comparative predictions that can serve as guidelines in biodiversity management.     

Multispecies coexistence of trees in tropical forests: spatial signals of topographic niche differentiation increase with environmental heterogeneity

Neutral and niche theories give contrasting explanations for the maintenance of tropical tree species diversity. Both have some empirical support, but methods to disentangle their effects have not yet been developed. We applied a statistical measure of spatial structure to data from 14 large tropical forest plots to test a prediction of niche theory that is incompatible with neutral theory: that species in heterogeneous environments should separate out in space according to their niche preferences. We chose plots across a range of topographic heterogeneity, and tested whether pairwise spatial associations among species were more variable in more heterogeneous sites. We found strong support for this prediction, based on a strong positive relationship between variance in the spatial structure of species pairs and topographic heterogeneity across sites. We interpret this pattern as evidence of pervasive niche differentiation, which increases in importance with increasing environmental heterogeneity.     

How do similarities in spatial distributions and interspecific associations affect the coexistence of Quercus species in the Baotianman National Nature Reserve,Henan, China

Congeneric species often have similar ecological characteristics and use similar resources. These similarities may make it easier for them to co‐occur in a similar habitat but may also lead to strong competitions that limit their coexistence. Hence, how do similarities in congeneric species affect their coexistence exactly? This study mainly used spatial point pattern analysis in two 1 hm² plots in the Baotianman National Nature Reserve, Henan, China, to compare the similarities in spatial distributions and interspecific associations of Quercus species. Results revealed that Quercus species were all aggregated under the complete spatial randomness null model, and aggregations were weaker under the heterogeneous Poisson process null model in each plot. The interspecific associations of Quercus species to non‐Quercus species were very similar in Plot 1. However, they can be either positive or negative in different plots between the co‐occurring Quercus species. The spatial distributions of congeneric species, interspecific associations with non‐Quercus species, neighborhood richness around species, and species diversity were all different between the two plots. We found that congeneric species did have some similarities, and the closely related congeneric species can positive or negative associate with each other in different plots. The co‐occurring congeneric species may have different survival strategies in different habitats. On the one hand, competition among congenerics may lead to differentiation in resource utilization. On the other hand, their similar interspecific associations can strengthen their competitive ability and promote local exclusion to noncongeneric species to obtain more living space. Our results provide new knowledge for us to better understand the coexistence mechanisms of species.     

The relationship between endophyte transition and plant species coexistence

The survival possibilities of terrestrial plant species are determined by their competitive abilities. One factor that affects competitive ability is the community of microorganisms that lives in association with the plants. Microorganisms affect the competitive dominance among plants by means of their metabolites. In this paper, we study the multiple plant species coexistence mediated by interactions with endophytes (fungi). The population dynamics are described by a revised lottery competition model for multiple plant species, each of which is divided into two classes: plants with endophytes (denoted EP) and plants without endophytes (NEP). The model includes the transition of seeds from EP to NEP. We show multiple species of plants cannot coexist in a steady state if this transition is density independent, but can coexist in a steady state if this transition is an increasing function of population density.     

Limited diffusive fluxes of substrate facilitate coexistence of two competing bacterial strains

Soils are known to support a great bacterial diversity down to the millimeter scale, but the mechanisms by which such a large diversity is sustained are largely unknown. A feature of unsaturated soils is that water usually forms thin, poorly-connected films, which limit solute diffusive fluxes. It has been proposed, but never unambiguously experimentally tested, that a low substrate diffusive flux would impact bacterial diversity, by promoting the coexistence between slow-growing bacteria and their potentially faster-growing competitors. We used a simple experimental system, based on a Petri dish and a perforated Teflon membrane to control diffusive fluxes of substrate (benzoate) whilst permitting direct observation of bacterial colonies. The system was inoculated with prescribed strains of Pseudomonas, whose growth was quantified by microscopic monitoring of the fluorescent proteins they produced. We observed that substrate diffusion limitation reduced the growth rate of the otherwise fast-growing Pseudomonas putida KT2440 strain. This strain out-competed Pseudomonas fluorescens F113 in liquid culture, but its competitive advantage was less marked on solid media, and even disappeared under conditions of low substrate diffusion. Low diffusive fluxes of substrate, characteristic of many unsaturated media (e.g. soils, food products), can thus promote bacterial coexistence in a competitive situation between two strains. This mechanism might therefore contribute to maintaining the noncompetitive diversity pattern observed in unsaturated soils.     

Density-dependent cooperation as a mechanism for persistence and coexistence

To overcome stress, such as resource limitation, an organism often needs to successfully mediate competition with other members of its own species. This may favor the evolution of defective traits that are harmful to the species population as a whole, and that may lead to its dilution or even to its extinction (the tragedy of the commons). Here, we show that this phenomenon can be circumvented by cooperation plasticity, in which an individual decides, based on environmental conditions, whether to cooperate or to defect. Specifically, we analyze the evolution of density-dependent cooperation. In our model, the population is spatially subdivided, periodically remixed, and comprises several species. We find that evolution pushes individuals to be more cooperative when their own species is at lower densities, and we show that not only could this cooperation prevent the tragedy of the commons, but it could also facilitate coexistence between many species that compete for the same resource.     

Abundance and spatial distribution of aphids and scales select for different life histories in their ladybird beetle predators

Abstract:  Life history parameters tend to differ between aphidophagous and coccidophagous ladybird beetles. It seems that the nature of prey, in particular the abundance, number and size of the colonies and their spatial distribution, may have been selected for the evolution of the life histories in these two groups of coccinellids, leading the aphidophagous ladybird beetles to develop at a fast pace and the coccidophagous beetles at a slower pace. To study the abundance, number and size of the colonies and the spatial distribution of aphid and coccid species, 100 sampling plots regularly spaced along four parallel transects were surveyed in the summer of 2004. At each sampling plot, species abundance, and the number and size of colonies of aphid and coccid species were recorded. Iwao's patchiness regression was used to assess the spatial distribution of aphids and coccids. From this study, it was found that coccids are much rarer than aphids but formed more colonies. Whereas aphids display a stonger tendency to crowding, aphid colonies are randomly distributed in space while coccid groups are aggregated. So, it seems that the abundance and spatial distribution of prey distribution may be factors selecting for the evolution of different life histories among aphidophagous and coccidophagous ladybird beetles.     

Heteromyopia and the spatial coexistence of similar competitors

Most spatial models of competing species assume symmetries in the spatial scales of dispersal and interactions. This makes analysis tractable, and has led to the conclusion that segregation of species in space does not promote coexistence. However, these symmetries leave parts of the parameter space uninvestigated. Using a moment‐approximation method, we present a spatial version of the Lotka–Volterra competition equations to investigate effects of removing symmetries in the distances over which individuals disperse and interact. Some spatial segregation of the species always comes about due to competition, and such segregation does not necessarily lead to coexistence. But, if interspecific competition occurs over shorter distances than intraspecific competition (heteromyopia), spatial segregation becomes strong enough to promote coexistence. Such coexistence is most likely when the species have similar dynamics, in contrast to the competition–colonization trade‐off that requires large competitive differences between species.     

Herbivore-induced coexistence of competing plant species

We study a series of spatially implicit lottery models in which two competing plant species, with and without defensive traits, are grazed by a herbivore in a homogeneous habitat. One species (palatable) has no defensive traits, while the other (defended) has defensive traits but suffers reduced reproduction as the result of an assumed trade-off. Not surprisingly, coexistence of these plants cannot occur when the herbivore density is very low (the palatable plant always wins) or very high (the defended plant wins). At intermediate densities, however, herbivory can mediate plant coexistence, even in a homogeneous environment. If the herbivore eats several plants per bite, and its forage-selection depends on the average palatability of the plants it eats, then palatable species in the immediate neighbourhood of defended plants may be more likely to persist (associational resistance) even at higher grazing pressure. If the herbivore shows a positive numerical response to the average palatability of the habitat as a whole, then both plant populations are stabilized and coexistence is promoted, because both species obtain a minority advantage through the negative feedback caused by herbivory. If the herbivore exhibits both of these traits, the system may have at most two non-trivial equilibria, one of which is stable and the other unstable. This means that coexistence in such a system is vulnerable to large fluctuations in herbivore density and identity, and this has implications for conservation in systems where large herbivores are managed to promote plant diversity.     

Local dispersal can facilitate coexistence in the presence of permanent spatial heterogeneity

Abstract In the presence of permanent spatial heterogeneity, local dispersal, especially short‐range dispersal, can facilitate coexistence by concentrating low‐density species in the areas where their rates of increase are higher. We present a framework for predicting the effects of local dispersal on coexistence for arbitrary forms of dispersal and arbitrary spatial patterns of environmental variation. Using the lottery model as an example, we find that local dispersal contributes to coexistence by enhancing the effects of environmental variation on scales longer than typical dispersal distances, which can be characterized solely by the variance of the dispersal kernel. Higher moments of the dispersal kernel are not important.