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Evolution of niche width and adaptive diversification.

Theoretical models suggest that resource competition can lead to the adaptive splitting of consumer populations into diverging lineages, that is, to adaptive diversification. In general, diversification is likely if consumers use only a narrow range of resources and thus have a small niche width. Here we use analytical and numerical methods to study the consequences for diversification if the niche width itself evolves. We found that the evolutionary outcome depends on the inherent costs or benefits of widening the niche. If widening the niche did not have costs in terms of overall resource uptake, then the consumer evolved a niche that was wide enough for disruptive selection on the niche position to vanish; adaptive diversification was no longer observed. However, if widening the niche was costly, then the niche widths remained relatively narrow, allowing for adaptive diversification in niche position. Adaptive diversification and speciation resulting from competition for a broadly distributed resource is thus likely if the niche width is fixed and relatively narrow or free to evolve but subject to costs. These results refine the conditions for adaptive diversification due to competition and formulate them in a way that might be more amenable for experimental investigations.

Adaptation, Physiological↗

Diversity studies of Salmonella incidents in some domestic livestock and their potential relevance as indicators of niche width.

This study attempts to determine whether or not livestock are becoming more susceptible to epidemics of salmonella infections by the analysis of published data on the annual number of reported infections in various animals. The number of incidents reported each year may be subject to a variety of biases due to temporal and geographical differences in reporting practices. This study analysed these reports by the calculation of diversity indices which are not subject to some of these potential biases. The relationship between the ecological concept of niche width and the diversity of species or types occupying that niche is discussed. The diversity of salmonella types reported in fowl has shown a highly significant decline over the 13-year period 1976-88. It is suggested that this declining diversity may be related to the declining niche width of the biotope available to this pathogen. Although speculative, this reduction in niche width could be related to a declining genetic diversity in the host animals or to an increasing intensification of animal husbandry.

Animals↗

Environmental Stresses Constrain Soil Microbial Community Functions by Regulating Deterministic Assembly and Niche Width.

Increasing evidence indicates that the loss of soil microbial α-diversity triggered by environmental stress negatively impacts microbial functions; however, the effects of microbial α-diversity on community functions under environmental stress are poorly understood. Here, we investigated the changes in bacterial and fungal α- diversity along gradients of five natural stressors (temperature, precipitation, plant diversity, soil organic C and pH) across 45 grasslands in China and evaluated their connection with microbial functional traits. By quantifying the five environmental stresses into an integrated stress index, we found that the bacterial and fungal α-diversity declined under high environmental stress across three soil layers (0-20 cm, 20-40 cm and 40-60 cm). Metagenomic-based analyses showed that the diversity of functional genes decreased along the stress gradients. High stress enhanced the abundance of genes associated with broad functional categories (e.g., glycolysis/gluconeogenesis, TCA cycle, DNA replication/repair and cell growth/death) but reduced the abundance of genes linked to specialised functional categories (e.g., C, N, S and methane metabolism). Phylogenetic null models and niche analyses indicated that stochastic assembly processes predominated in high-diversity communities, in which bacterial and fungal taxa had a narrow ecological niche. However, in low-diversity communities, deterministic assembly processes were dominant, and taxa had wide niches, correlating with the reduction in gene abundance observed for broad and specialised functional categories. Given the essential role of the microbiome in regulating ecosystem functions, our findings suggest that low-diversity-induced deterministic community assembly processes and a wide niche under high environmental stress may regulate microbial functions. These findings emphasise the ecological mechanisms through which microbial biodiversity regulates terrestrial ecosystem functioning.

Soil Microbiology↗

Intraspecific competition favours niche width expansion in Drosophila melanogaster.

Ecologists have proposed that when interspecific competition is reduced, competition within a species becomes a potent evolutionary force leading to rapid diversification. This view reflects the observation that populations invading species-poor communities frequently evolve broader niches. Niche expansion can be associated with an increase in phenotypic variance (known as character release), with the evolution of polymorphisms, or with divergence into many species using distinct resources (adaptive radiation). The relationship between intraspecific competition and diversification is known from theory, and has been used as the foundation for some models of speciation. However, there has been little empirical proof that niches evolve in response to intraspecific competition. To test this hypothesis, I introduced cadmium-intolerant Drosophila melanogaster populations to environments containing both cadmium-free and cadmium-laced resources. Here I show that populations experiencing high competition adapted to cadmium more rapidly than low competition populations. This provides experimental confirmation that competition in a population can drive niche expansion onto new resources for which competition is less severe.

Adaptation, Physiological↗

[Quantitative characteristics of degenerative succession in Festuca sinensis sowing grassland in the alpine pastoral area].

Quantitative characteristics of degenerative succession in Festuca sinensis sowing grassland in the alpine pastoral area were studied. The results showed that with the succession developing, number of species, richness index, diversity index, and evenness index were all increased, but ecological dominance was decreased. Analysis on important values, niche width, and niche overlaps proved that some herb species such as Ajania tenuifolia and Polygomum sibiricum were pioneer plants invading sowing grassland in the succession process. The pioneer plants had wider niche widths than other species, and their important values increased with the succession developing. Niche overlaps between Festuca sinensis and other species were smaller than that between two latifoliate herbs species in most case. The invasion of latifoliate herbs was one of the most important factors, which made the sowing grassland more degenerative.

Cold Climate↗

Niche packing and coevolution in competition communities.

A coevolutionary model of species packing is developed that allows evolutionary adjustment in both niche position and within-phenotype niche width of one-three competing species. The environment is specified as a single resource dimension x and availability of resources along x is given by a gaussian curve that has parameters x[unk](R) and sigma(R). The model predicts that, for S species, the ratio of optimal niche width w[unk] to sigma(R) is roughly independent of sigma(R) and can be approximated by 1/S when the competitors are completely resource limited. Niche separation (d[unk]/w[unk]) increases only moderately with increases in resource diversity sigma(R) and is greater for two than for three competing species. To the extent that the competitors are not completely resource limited, both coevolutionary niche separation and niche width decrease. Many of the general trends in niche width and niche separation predicted by this coevolutionary model parallel those from optimal foraging theory and limiting similarity models of community structure. The coevolutionary model stands out, however, in the singularly high values predicted for niche separation, making coevolved communities highly invasible. Hence, the theory suggests, as some empirical evidence indicates, that coevolved competition communities can only eixst as such on remote islands or in other habitats that might be free from invasion by outside species.

Journal Article↗

Population dynamics of phytophagous and predatory mites (Acari: Tetranychidae, Eriophyidae, Phytoseiidae) on bamboo plants in Fujian, China.

The seasonal cycle and population dynamics of Schizotetranychus nanjingensis Ma and Yuan, Aponychus corpuzae Rimando (Tetranychidae), Aculus hambusae Kuang (Eriophyidae) and their natural enemy Tvphlodromus bambusae Ehara (Phytoseiidae) were studied during 1996-1998 in moso bamboo forests in Nanping, Fujian, China. Damage to bamboo leaves was often caused by mixed populations of the three phytophagous species, which displayed different seasonal dynamics: S. nanjingensis fed actively and reproduced in May and from late July to late October (aestivating from June to mid July), Ap. corpuzae was active and reproduced from mid July to late November, and Ac. bambusae from July to next February. During Spring S. nanjingensis was the dominant species with the greatest niche width, but in Summer Ac. bambusae and Ap. corpuzae became dominant and had greater niche widths. In Autumn, Ac. bambusae decreased but both S. nanjingensis and Ap. corpuzae mites increased and the latter became dominant. In winter all species decreased sharply in number. The two spider mite species (S. nanjingensis and Ap. corpuzae) had high niche overlaps in all four seasons. Niche overlaps between the eriophyid Ac. bambusae and the two spider mites were similarly high except during the spring when Ac. bambusae was absent. The predatory mite (T. bambusae) had higher niche overlap with Ap. corpuzae than with others during Autumn and Winter, but during Spring and Summer niche overlap was higher with S. nanjingensis.

Animals↗

Interspecific interactions between Acanthocephala in the intestine of brown trout: are they more frequent in Ireland?

The aim of this paper was to test the hypothesis that when the 2 species of Acanthocephalan Pomphorhynchus laevis and Acanthocephalus clavula are found concurrently within the intestine of brown trout under field conditions, they have the potential to interact negatively. Evidence has shown that Acanthocephala are more likely to exhibit negative interactions with their own and other species, under both field and experimental conditions. Furthermore, the likelihood of these interactions is increased in Ireland because of the absence of certain definitive hosts and the fact that concurrent infections by two or more species of Acanthocephala are more commonly observed in fish. Data collected from wild and stocked brown trout and from 2 lakes provided an opportunity to compare the 2 potentially interacting helminth species in their fundamental and realized niche and several pieces of convincing evidence are provided here to support the hypothesis. A significant negative association between the numbers of each species found in individual fish was reported and this was consistent for both wild and stocked trout. Furthermore, an analysis of the proportions of low, moderate and high intensity infections in single and concurrent infections revealed a significant reduction in increasing intensities in concurrent infections compared to single infections. Finally, strikingly different patterns of niche inhabitation were observed, particularly for P. laevis in the presence of A. clavula in wild trout. Results from the niche width analysis also support the observations on average position in single and concurrent infections. The niche width of P. laevis when it co-occurred with A. clavula decreased markedly in high intensity infections compared to low intensity infections.

Acanthocephala↗

Culturability as an indicator of succession in microbial communities.

Successional theory predicts that opportunistic species with high investment of energy in reproduction and wide niche width will be replaced by equilibrium species with relatively higher investment of energy in maintenance and narrower niche width as communities develop. Since the ability to rapidly grow into a detectable colony on nonselective agar medium could be considered as characteristic of opportunistic types of bacteria, the percentage of culturable cells may be an indicator of successional state in microbial communities. The ratios of culturable cells (colony forming units on R2A agar) to total cells (acridine orange direct microscopic counts) and culturable cells to active cells (reduction of 5-cyano-2,3-ditolyl tetrazolium chloride) were measured over time in two types of laboratory microcosms (the rhizosphere of hydroponically grown wheat and aerobic, continuously stirred tank reactors containing plant biomass) to determine the effectiveness of culturabilty as an index of successional state. The culturable cell:total cell ratio in the rhizosphere decreased from approximately 0.25 to less than 0.05 during the first 30-50 days of plant growth, and from 0.65 to 0.14 during the first 7 days of operation of the bioreactor. The culturable cell:active cell ratio followed similar trends, but the values were consistently greater than the culturable cell:total cell ratio, and even exceeded I in early samples. Follow-up studies used a cultivation-independent method, terminal restriction fragment length polymorphisms (TRFLP) from whole community DNA, to assess community structure. The number of TRFLP peaks increased with time, while the number of culturable types did not, indicating that the general decrease in culturability is associated with a shift in community structure. The ratio of respired to assimilated C-14-labeled amino acids increased with the age of rhizosphere communities, supporting the hypothesis that a shift in resource allocation from growth to maintenance occurs with time. Results from this work indicate that the percentage of culturable cells may be a useful method for assessing the successional state of microbial communities.

Amino Acids↗

Genetic diversity and its ecological correlates in nature: comparisons between subterranean, fossorial, and aboveground small mammals.

We have reanalyzed the levels of observed heterozygosity based on old and new material of small mammals in attempt to retest the environmental theory of genetic diversity or the niche-width variation hypothesis. This study involves old and new material of 184 small mammalian species both insectivores and rodents; 111 living aboveground and 73 species living partly (fossorial, N = 28) or totally (subterranean, N = 45) underground. The results indicated that (i) in general, insectivores harbor significantly lower levels of heterozygosity than rodents, and (ii) both overall, and in insectivores and rodents separately, the narrow-niche fossorial and subterranean species displayed significantly lower levels of observed heterozygosity than small mammalian species living aboveground. This is true in almost all parametric and nonparametric analyses. Subterranean and fossorial small mammals are more homozygous in protein diversity, primarily isozymes, compared with aboveground species. This pattern corroborates the environmental theory or niche-width variation hypothesis of genetic diversity.

Animals↗

Niche restriction in parasites: proximate and ultimate causes.

Hutchinson's (1957) definition of an ecological niche as a multidimensional hypervolume determined by a number of physical and biotic variables is adopted. The number of niche dimensions is very great, but as a working hypothesis it is assumed that a few are sufficient to characterize the niche of a parasite species to a high degree of accuracy. They are host species, microhabitat(s), macrohabitat(s), geographical range, sex and age of host, season, food and hyperparasites. Methods to measure niche width, in particular specificity indices, are discussed, and some examples of niche restriction are described. Proximate and ultimate causes of niche restriction are discussed, mainly using marine parasites as examples. Among proximate causes of one niche dimension, host specificity, are ecological factors restricting exposure to infection to certain host species; host-specific chemical factors that induce hatching, direct infective stages to a host and bring about settlement of a parasite; factors that lead to mortality in or on the wrong host; morphological adaptations that guarantee survival in or on the 'correct' host; and availability of suitable hosts. Many factors are likely to be responsible for microhabitat specificity, but have been little studied, except for some physiological and morphological adaptations to particular microhabitats. Macrohabitats and geographical range may be determined by the distribution of intermediate hosts and certain food items, and by a variety of chemical and physical factors. Hosts of different sexes may differ in feeding habits and the composition of the skin, and thus acquire parasites differentially. Hosts of different age may be differentially infected due to accumulation of parasites with age, loss of parasites due to developing resistance (or immunity), and different size and feeding habits. Among ultimate causes of niche restriction and segregation are avoidance of competition, predation and hyperparasites; facilitation of mating; reinforcement of reproductive barriers; and adaptations to environmental complexity. Few studies permit a decision on which factor or factors are responsible in particular cases. Interspecific competition may play a greater role in helminth communities of some host groups than of others, but it seems that, overall, its role has been exaggerated at least for marine parasites. Some 'classical' examples of microhabitat segregation explained by interspecific competition can also be explained by reinforcement of reproductive barriers. There is evidence for the importance of facilitation of mating in microhabitat restriction, and the availability of many vacant niches indicates that competition, overall, is not of great importance.

Animals↗

Resource variation and the structure of British bird communities.

Data on the foraging microhabitats of British birds are reanalyzed with the aim of understanding how fluctuations in resource abundance affect niche relationships and community structure. At Marley Wood, overlap in the foraging sites of resident bird species increased during the late spring and summer and decreased during the fall and winter. Among bird species coexisting in the pine forests at Thetford Chase, spatial overlap and spatial niche widths were positively correlated with food abundance over a 4-year period. These results suggest that in variable environments similarity in spatial niches is an inverse function of the intensity of competition for food. As food supplies drop, consumer species must apparently occupy increasingly different foraging areas in order to coexist. In less variable environments, however, resource stability may allow finer partitioning of the available foraging space and greater spatial overlap within a given foraging area. The results of this paper also suggest a reinterpretation of MacArthur's study of resource partitioning among warbler species in the boreal forests of New England. Rather than providing an instance of niche segregation in order to avoid intense competition, MacArthur's warblers may actually represent another example of increased spatial similarity when food resources are abundant and competition is reduced.

Journal Article↗

The ecology of individuals: incidence and implications of individual specialization.

Most empirical and theoretical studies of resource use and population dynamics treat conspecific individuals as ecologically equivalent. This simplification is only justified if interindividual niche variation is rare, weak, or has a trivial effect on ecological processes. This article reviews the incidence, degree, causes, and implications of individual-level niche variation to challenge these simplifications. Evidence for individual specialization is available for 93 species distributed across a broad range of taxonomic groups. Although few studies have quantified the degree to which individuals are specialized relative to their population, between-individual variation can sometimes comprise the majority of the population's niche width. The degree of individual specialization varies widely among species and among populations, reflecting a diverse array of physiological, behavioral, and ecological mechanisms that can generate intrapopulation variation. Finally, individual specialization has potentially important ecological, evolutionary, and conservation implications. Theory suggests that niche variation facilitates frequency-dependent interactions that can profoundly affect the population's stability, the amount of intraspecific competition, fitness-function shapes, and the population's capacity to diversify and speciate rapidly. Our collection of case studies suggests that individual specialization is a widespread but underappreciated phenomenon that poses many important but unanswered questions.

Animals↗

Constraints in temperature adaptation reinforce differences in thermal niche between mesophilic and psychrotolerant Bacillus cereus group species.

Experimental evolution has demonstrated that mesophilic microbes readily adapt to increases in temperature. However, many microbes are psychrotolerant and resistant to cold, which is associated with physiological specializations, suggesting constraints in thermal adaptation. We hypothesized that constraints would limit adaption differently in a mesophilic species (Bacillus thuringiensis) compared with its psychrotolerant relative B. mycoides-with adaptation at cooler temperatures and adaptation at higher temperatures being constrained in each species, respectively. To test this hypothesis, we imposed 140 generations of selection at temperatures at and below the optimum for productivity for both species. The fitness and thermal performance of evolved bacteria showed ancestral thermal niche plays a role in thermal adaptation over this time scale, in support of our hypothesis of adaptive constraints. Temperature-dependent trade-offs appeared common in B. mycoides, with fitness gains associated with decreases in operational niche width; fitness gains at one temperature caused a decrease in the range of temperatures that the bacterium showed appreciable growth. Genome resequencing showed that variation in mutation supply and selection strength could not explain temperature-dependent responses to selection. Importantly, metabolic theory only held true for mesophilic B. thuringiensis, showing abundant but less studied psychrotolerant species could follow different adaptive trajectories.

Bacillus thuringiensis↗

Parasite communities of the Salzhaff (Northwest Mecklenburg, Baltic Sea). I. Structure and dynamics of communities of littoral fish, especially small-sized fish.

The parasites of ten fish species, including four Gobiidae, three Gasterosteidae, two Syngnathidae, and one Zoarcidae from the Salzhaff region, Northwest Mecklenburg, Baltic Sea, were investigated in 1995 and 1996. As many as 36 parasite species, represented by diverse groups of helminths and protozoans as well as annelids and copepods, are found during 4 seasons in these hosts. By far the most abundant group is represented by digeneans, comprising 15 species, followed by 7 cestodans and 6 nematodes. With regard to component communities, 8 host-parasite combinations are core and secondary species with more than 40% prevalence in which generalists such as the digeneans Podocotyle atomon and Cryptocotyle concavum (in 3 hosts). C. lingua, Diplostomum spathaceum, and Acanthostomum balthicum, and the nematode Hysterothylacium are involved. Also, specialists such as Aphalloides timmi in Pomatoschistus microps as well as Magnibursatus caudofilamentosa and Thersitina gasterostei in Gasterosteus aculeatus attain high levels of prevalence. A comparison of different investigations reveals greater prevalence of allogenic and autogenic parasite species with 3 host cycles in the Rerik-Riff (free Baltic) and higher levels of prevalence of autogenic parasite species with 1 or 2 host cycles in the entire Salzhaff. The component communities of gobies from Dahmeshöved, Lübeck Bight, attain generally lower degrees of prevalence than those of the Salzhaff region. The infracommunities consist mostly of 1-3 parasite species per host specimen; this value is surpassed on occasion in P. microps (maximum 7 species) and in G. aculeatus (maximum 9 species, which may compete for 5 microhabitats in a host specimen). In this context the theory of empty niches propagated by some parasitologists is critically discussed and substituted by the assumption of variable niche widths. The seasonality of the more abundant parasites is either unclear--as in the case of C. concavum--or evident--as in the case of P. atomon, which prevail in early spring and summer, or A. timmi, which dominate in late summer, as do M. caudofilamentosa, which is absent in spring. The main causes of the infestation of fish hosts may be their ages and the availability of parasites due to the presence of intermediate hosts.

Animals↗

Multi-species outcomes in a common model of sympatric speciation.

While models of sympatric speciation are motivated in part by multi-species adaptive radiations such as the Cameroon crater lake cichlids, existing models have focused on bifurcation into a single pair of daughter species. This paper shows that a familiar model of sympatric speciation, driven by intraspecific competition and assortative mating based on ecological characters values, can yield multiple daughter species if individual niche widths are sufficiently restricted. Surprisingly, the multi-species outcome is not produced by successive bifurcation events, but by simultaneous divergence resulting in a hard polytomy. This result is sensitive to a number of assumptions, whose violation may prevent speciation. In some cases when speciation fails, the population instead ends in a state that closely resembles incipient species pairs, with an ecological polymorphism and partial reproductive isolation. However, this polymorphism is stable and does not lead to complete reproductive isolation, suggesting that empirical cases of incipient species pairs may not always end in speciation.

Animals↗

Genetic polymorphisms in subterranean mammals (Spalax ehrenbergi superspecies) in the near east revisited: patterns and theory.

Allozyme diversity in the superspecies Spalax ehrenbergi has been revisited by studying 36 gene loci in 241 subterranean mole rats from 22 populations and nine chromosomal species, four from Turkey (2n = 52E (east), 52W (west), 56 and 58), four from Israel (2n = 52, 54, 58 and 60), and one from Egypt (2n = 60). The following results were indicated. (1) Genetic patterns: 11 of the 36 loci analysed (30.5 per cent) were monomorphic across the range, fifteen (41.7 per cent) were weakly polymorphic and the remaining 10 loci (27.8 per cent) were strongly polymorphic. (2) Heterozygosity: the average H was 0.051, range 0.00-0.098. In Israel, H increased with aridity and climatic unpredictability towards the northern Negev Desert, and was remarkably high in small steppic semi-isolates and desert isolates. (3) Species discrimination: some of the S. ehrenbergi species can be discriminated qualitatively. (4) Genetic distances (D): between species these values averaged 0.077, range 0.001-0.269, with the highest D between the ancestor Turkish and descendant Israeli and Egyptian species. The phylogenetic tree supports the Turkish origin of the Israeli Spalax ehrenbergi species, and the recent speciation of the Egyptian Spalax. (5) Genetic diversity is mostly (58 per cent) within populations. (6) Allozyme correlates: allozyme diversity was significantly correlated with the external physical (both climatic and edaphic) and biotic (parasite infection and plant cover) environment. (7) Spatial autocorrelation of allozyme frequencies suggests that migration is not influential. (8) Gametic phase disequilibria were significant in four out of five species tested, and were associated with climatic and edaphic factors. These results support the environmental selection hypothesis of genetic diversity including the niche-width variation hypothesis in space and time. Natural selection appears to play a major role in genetic differentiation of proteins in adaptive radiation and speciation.

Animals↗

Niche tradeoffs, neutrality, and community structure: a stochastic theory of resource competition, invasion, and community assembly.

Stochastic niche theory resolves many of the differences between neutral theory and classical tradeoff-based niche theories of resource competition and community structure. In stochastic niche theory, invading species become established only if propagules can survive stochastic mortality while growing to maturity on the resources left unconsumed by established species. The theory makes three predictions about community structure. First, stochastic niche assembly creates communities in which species dominate approximately equally wide "slices" of the habitat's spatial heterogeneity. These niche widths generate realistic distributions of species relative abundances for which, contrary to neutral theory but consistent with numerous observations, there are strong correlations among species traits, species abundances, and environmental conditions. Second, slight decreases in resource levels are predicted to cause large decreases in the probability that a propagule would survive to be an adult. These decreases cause local diversity to be limited by the inhibitory effects of resource use by established species on the establishment (recruitment) of potential invaders. If resource pulses or disturbance allowed invaders to overcome this recruitment limitation, many more species could indefinitely coexist. Third, the low invasibility of high diversity communities is predicted to result not from diversity per se, but from the uniformly low levels of resources that occur in high-diversity communities created by stochastic competitive assembly. This prediction provides a potential solution to the invasion paradox, which is the tendency for highly diverse regions to be more heavily invaded.

Computer Simulation↗