Host-parasite coevolution: comparative evidence for covariation of life history traits in primates and oxyurid parasites.

The environmental factors that drive the evolution of parasite life histories are mostly unknown. Given that hosts provide the principal environmental features parasites have to deal with, and given that these features (such as resource availability and immune responses) are well characterized by the life history of the host, we may expect natural selection to result in covariation between parasite and host life histories. Moreover, some parasites show a high degree of host specificity, and cladistic analyses have shown that host and parasite phylogenies can be highly congruent. These considerations suggest that parasite and host life histories may covary. The central argument in the theory of life history evolution concerns the existence of trade-offs between traits. For parasitic nematodes it has been shown that larger body sizes induce higher fecundity, but this is achieved at the expense of delayed maturity. As high adult mortality would select for reduced age at maturity, the selective benefit of increased fecundity is expressed only if adult mortality is low. Parasite adult mortality may depend on a number of factors, including host longevity. Here we tested the hypothesis concerning the positive covariation between parasite body size (which reflects parasite longevity) and host longevity. To achieve this goal, we used the association between the pinworms (Oxyuridae, Nematoda) and their primate hosts. Oxyurids are highly host specific and are supposed to be involved in a coevolutionary process with their hosts. We found that female parasite body length was positively correlated with host longevity after correcting for phylogeny and host body mass. Conversely, male parasite body length and host longevity were not correlated. These results confirm that host longevity may represent a constraint on the evolution of body size in oxyurids, at least in females. The discrepancy between female and male oxyurids is likely to depend on the particular mode of reproduction of this taxon (haplodiploidy), which should result in weak (or even null) selection pressures to an increase of body size in males.     

Genetic variation and covariation among life history traits in populations of Acanthoscelides obtectus maintained on different hosts

We explored the extent to which two populations of Acanthoscelides obtectus (Coleoptera: Bruchidae), maintained in the laboratory for about 60 generations on common bean (Phaseolus vulgaris) and chickpea (Cicer arietinum) seeds, differ genetically in life history traits. Using a half-sib breeding design, we screened for genetic variation in oviposition preference for two hosts within both populations, and examined whether this variation is correlated with weevil pre-adult (egg-to-adult viability and egg-to-adult developmental time) and adult (female fecundity and longevity) fitness components on these two hosts. Significant differences between the Phaseolus and Cicer population were detected in viability, developmental time and oviposition preference, but not in female fecundity and longevity. The absence of a significant population × host interaction in all traits, except for developmental time in males, indicates that the Cicer population has not attained the status of host race. The estimated narrow-sense heritabilities within population within host plant were in the low-to-moderate level (range: 1%–79%), with an overall average of about 24%. Within the population maintained on the chickpeas we detected significant genetic correlations between female developmental time on chickpea seeds and preference for chickpeas. In the population maintained on the primary host of this weevil species (the Phaseolus population) we found no significant preference/performance genetic correlations. Data are also presented on the genetic correlations of preference and performance across host seeds within both populations. These correlations are found to be positive for all traits in both populations, indicating the absence of trade-offs in either performance or behavioural response that might impose selection for host specialization within populations.     

Climate envelope, life history traits and the resilience of birds facing global change

Few studies have examined how life history traits and the climate envelope influence the ability of species to respond to climate change and habitat degradation. In this study, we test whether 18 species-specific variables, related to the climate envelope, ecological envelope and life history, could predict recent population trends (over 17 years) of 71 common breeding bird species in France. Habitat specialists were declining at a much higher rate than generalists, a sign that habitat quality is decreasing globally. The lower the thermal maximum (temperature at the hot edge of the climate envelope), the more negative are the population trends and the less tolerant these species are climate warming, regardless of the thermal range over which these species occur. The life history trait 'the number of broods per year' was positively related to recent trends, suggesting that single-brooded species might be more sensitive to advances in food peak due to climate change, as it increases the risk of mistiming their single-breeding event. Annual fecundity explained long-term declines, as it is a good proxy for most other demographic rates, with shorter-lived species being more sensitive to global change: individuals of species with higher fecundity might have too short a life to learn to adapt to directional changes in their environment. Finally, there was evidence that natal dispersal was a predictor of recent trends, with species with high natal dispersal experiencing smaller population declines than species with low natal dispersal. This is expected if the higher the natal dispersal, the larger the ability to shift spatially when facing changes in local habitat or climate, in order to track optimal conditions and adapt to global change. Identifying decline-promoting factors allow us to infer mechanisms responsible for observed declines in wild bird populations facing global change, and by doing so allow for a more pre-emptive approach to conservation planning.     

Perinatal life history traits in New World monkeys

Gestation length, neonatal and maternal body weight, and neonatal and adult brain weight data were collected for New World monkeys in an attempt to establish typical patterns of perinatal life history. This study attempts to illuminate the most accurate values from the available data, which suggest that gestation length and prenatal growth rate are broadly conserved in relation to maternal size in New World monkeys. Exceptions to the patterns evident in the data point to derivations in life history strategies. In particular, this study suggests that the extended gestation length of callitrichines is a function of minimum viable neonate size and not exclusively energy minimization associated with simultaneous lactation. Cebus is shown to undergo more postnatal brain growth relative to other New World monkeys, but not as much as previously believed. Alouatta is shown to be relatively small brained at birth as well as in adulthood. Saimiri is shown to present the most unusual package of perinatal life history traits, in which precocial neonates are gestated for a relatively long time and at a slightly faster growth rate than is typical for New World monkeys. © 1996 Wiley-Liss, Inc.     

Environmentally induced clonal reproduction and life history traits of the liana Machaerium cuspidatum in an Amazonian rain forest,Ecuador

The population structure of the canopy liana Machaerium cuspidatum Kuhlm. & Hoehne (Fabaceae) was studied in ten transects covering a total of 2.5 ha of tropical rain forest in Yasuní National Park, eastern Ecuador. The aim was to investigate how trellis availability, topographic position and light availability affected the population density and structure of the most abundant liana species in the area. The environmental variation affected plants in different size classes differently. Large plants (diameter 1 cm) were almost exclusively found in areas with low canopy and dense undergrowth. These areas had the highest density of suitable host plants. Seedling-sized plants (height < 30 cm) and saplings (height 30 cm but diameter < 1 cm) occurred more frequently and withhigher density in steep upland areas than in the floodplains, presumably due to elevated seedling mortality in the periodically flooded areas. Seedling-sized plants and saplings did not grow in areas with elevated light levels, and although plants of all sizes were highly clumped, this clumping rarely coincided with patches with abundant canopy gaps. Machaerium cuspidatum reproduces clonally when branches re-root and break off from the parentplant. The density of clonally produced plants was high in areas where the density of sexually produced plants was low, whereas the total density varied little among habitat types. A higher proportion of the seedling-sized plants were of sexual origin in the upland areas. An environmentally induced increase in the production of independent ramets enables M. cuspidatum to persist in the floodplains. This shows that plants may expand theirrealised niches to habitats with a low potential for seedling survival if high seedling mortality can be compensated for by increased clonal reproduction.     

Genetic architecture of life history traits and environment-specific trade-offs

Life history theory predicts the evolution of trait combinations that enhance fitness, and the occurrence of trade-offs depends in part on the magnitude of variation in growth rate or acquisition. Using recombinant inbred lines, we examined the genetic architecture of age and size at reproduction across abiotic conditions encountered by cultivars and naturalized populations of Brassica rapa. We found that genotypes are plastic to seasonal setting, such that reproduction was accelerated under conditions encountered by summer annual populations and genetic variances for age at reproduction varied across simulated seasonal settings. Using an acquisition-allocation model, we predicted the likelihood of trade-offs. Consistent with predicted relationships, we observed a trade-off where early maturity is associated with small size at maturity under simulated summer and fall annual conditions but not under winter annual conditions. The trade-off in the summer annual setting was observed despite significant genotypic variation in growth rate, which is often expected to decouple age and size at reproduction because rapidly growing genotypes could mature early and attain a larger size relative to slowly growing genotypes that mature later. The absence of a trade-off in the winter setting is presumably attributable to the absence of genotypic differences in age at reproduction. We observed QTL for age at reproduction that jointly regulated size at reproduction in both the summer and fall annual settings, but these QTL were environment-specific (i.e. different QTL contributed to the trade-off in the fall vs. summer annual settings). Thus, at least some of the genetic mechanisms underlying observed trade-offs differed across environments.     

Ecological immunology: life history trade-offs and immune defense in birds

There has been considerable recent interest in the effects oflife-history decisions on immunocompetence in birds. If immunocompetenceis limited by available resources, then trade-offs between investmentin life-history components and investment in immunocompetencecould be important in determining optimal life-history traits.For this to be true: (1) immunocompetence must be limited byresources, (2) investment in life-history components must benegatively correlated with immunocompetence, and (3) immunocompetencemust be positively correlated with fitness. To gather such empiricaldata, ecologists need to be able to measure immunocompetence.We review techniques used to measure immunocompetence and howthey are applied by ecologists. We also consider the componentsof the immune system that constitute immunocompetence and evaluatethe possible consequences of measuring immunocompetence in differentways. We then review the empirical evidence for life-historytrade-offs involving immune defense. We conclude that thereis some evidence suggesting that immunocompetence is limitedby resources and that investment in certain life-history componentsreduces immunocompetence. However, the evidence that immunocompetenceis related to fitness is circumstantial at present, althoughconsistent with the hypothesis that immunocompetence and fitnessare positively correlated. We argue that future work needs toexamine the fitness effects of variation in immunocompetenceand suggest that artificial selection experiments offer a potentiallyimportant tool for addressing this issue.     

Multiple growth-correlated life history traits estimated simultaneously in individuals

We present a new methodology to estimate rates of energy acquisition, maintenance, reproductive investment and the onset of maturation (four-trait estimation) by fitting an energy allocation model to individual growth trajectories. The accuracy and precision of the method is evaluated on simulated growth trajectories. In the deterministic case, all life history parameters are well estimated with negligible bias over realistic parameter ranges. Adding environmental variability reduces precision, causes the maintenance and reproductive investment to be confounded with a negative error correlation, and tends, if strong, to result in an underestimation of the energy acquisition and maintenance and an overestimation of the age and size at the onset of maturation. Assuming a priori incorrect allometric scaling exponents also leads to a general but fairly predictable bias. To avoid confounding in applications we propose to assume a constant maintenance (three-trait estimation), which can be obtained by fitting reproductive investment simultaneously to size at age on population data. The results become qualitatively more robust but the improvement of the estimate of the onset of maturation is not significant. When applied to growth curves back-calculated from otoliths of female North Sea plaice Pleuronectes platessa , the four-trait and three-trait estimation produced estimates for the onset of maturation very similar to those obtained by direct observation. The correlations between life-history traits match expectations. We discuss the potential of the methodology in studies of the ecology and evolution of life history parameters in wild populations.     

Variation in life history and genetic traits of Hawaiian mosquitofish populations

Mosquitofish (Gambusia affinis) were collected from 17 reservoirs on three islands in Hawaii, USA. Genetic and life history traits for adult females from these populations were used to evaluate hypotheses concerning short-term evolutionary divergence of populations recently established from a common ancestral source. The effects of founder events and drift on genetic variability and population differentiation were also examined. Significant differences in life history characteristics, allele frequencies, and multi-locus heterozygosities (H) were found among fish populations collected from different reservoirs and between reservoirs classified as stable or fluctuating on the basis of temporal fluctuation in water level. Females from stable reservoirs exhibited greater standard length (35.1 vs 32.8 mm), lower fecundity (11.9 vs 15.2 embryos), lower reproductive allocation (18.2% vs 22.8%), but larger mean embryo size (1.95 vs 1.67 mg) than females from fluctuating reservoirs. Consistency in means among replicates of each reservoir class and concordance in direction and magnitude of differences reported here and results of sampling conducted from these same locations 10 years previously (Stearns, 1983a) suggest that ecological factors intrinsic to these two environments are important in determing population life history traits. Females from stable reservoirs exhibited lower heterozygosity than females from fluctuating reservoirs (0.134 vs 0.158, respectively). Levels and direction od differences in heterozygosity, the high proportion of polymorphic loci and lack of fixation of alternative alleles argue against a purely stochastic explanation for genetic and life history variation among reservoir populations. Levels of genetic variability and interpopulation differentiation were similar to those observed in mainland populations of this species. A high proportion of the genetic diversity was apportioned between populations and within populations due to differences between juveniles and adults. Significant genotypic differences between adult and juvenile age classes suggest that the genetic divergence of local populations may occur over short periods of time.     

Coevolution of life history traits and morphology in female subterranean amphipods

Subterranean species show a distinct morphology, yet the adaptive significance of some traits, like body size and shape, is poorly understood and cannot be explained solely by distinct environmental conditions (darkness, less food). We predicted that in females some morphological changes may have co‐evolved with life history traits, and that co‐evolving life history traits provide at least part of the explanation for evolutionary changes of morphology. Using museum material we tested this prediction on the subterranean amphipod genus Niphargus. We studied six species found in springs and eight species found in cave lakes. We treated them as two ecologically distinct groups, and the major ecological differences between them were the availability of nutrients and the water currents. Cave species were found to be larger and stouter (as inferred from the shape of coxal plates, which are part of the marsupium), they had larger eggs and lower reproductive effort per brood, whereas the egg number and brood volume if corrected for the body size were not different. Using phylogenetic independent contrasts, we found a positive correlation between body shape and egg volume, a positive correlation between body size and egg volume, and a negative correlation between body size and reproductive effort per brood. We tentatively conclude that evolutions of morphology and life histories are functionally connected and that co‐evolving traits contribute to overall selective regime.     

Sexual differences in larval life history traits of acanthocephalan cystacanths

Sexual differences in life history traits, such as size dimorphism, presumably arise via sexual selection and are most readily observed in adults. For complex life-cycle parasites, however, sexual selection may also have consequences for larval traits, e.g., growth in intermediate hosts. Two acanthocephalan species (Acanthocephalus lucii and Echinorhynchus borealis) were studied to determine, whether larval life histories differ between males and females. The size of female A. lucii cystacanths had a much stronger relationship with intermediate host size than males, suggesting females invest more in growth and are consequently more limited by resources. No relationship between host size and cystacanth size was observed for E. borealis. For both species, female cystacanths survived longer in a culture medium composed entirely of salts, which could suggest that females have greater energy reserves than males. A comparative analysis across acanthocephalan species indicated that sexual size dimorphism at the adult stage correlates with cystacanth dimorphism. However, the relationship was not isometric; cystacanths do not reach the same level of sexual dimorphism as adults, possibly due to resource constraints. Our results suggest that larval life histories diverge between males and females in some acanthocephalans, and this is seemingly a consequence of sexual selection acting on adults.     

Dietary effects on life history traits of riverine Bosmina

1. We compared growth, reproduction and life history characteristics of Bosmina raised on Ohio River seston versus a unialgal culture (Scenedesmus acutus), to assess potential nutritional constraints experienced by riverine populations. 2. Bosmina grew well in both treatments during their juvenile stage. Analysis of variance showed that Bosmina growth prior to the start of reproduction did not differ significantly between the treatments. After the onset of reproduction Bosmina fed on Scenedesmus grew faster and exhibited higher fecundity than their counterparts fed river seston. 3. Significant quantities of triacylglycerol (visible lipid droplets) were gradually accumulated in Bosmina fed on Scenedesmus. Visual lipid ovary indices were positively correlated with adult growth rate and fecundity and negatively correlated with longevity. 4. Biochemical analysis showed that Scenedesmus had significantly greater total ω3, ω6 and polyunsaturated fatty acids (PUFA) whereas river seston had more eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA). Thus, in contrast to Daphnia, EPA and DHA do not appear to influence Bosmina growth or reproductive performance, but linolenic acid or total PUFA may be important. 5. Comparatively weak performance by Bosmina feeding on river seston suggest that nutritional constraints may be important despite high particulate organic carbon and low C : N and C : P of river seston.     

Seedling herbivory, community composition and plant life history traits

Despite the voluminous literature documenting plant-animal interactions, it is only recently that ecologists have begun to focus upon the importance of seedling herbivory in plant communities. This review aims to synthesize our current understanding of the effect that selective seedling removal by herbivores has in shaping post-disturbance plant community structure. In order to do this I describe how individual seedling and plant community characteristics influence the likelihood of seedling herbivory. I argue that seedling palatability, size and morphology, together with the frequency, distribution and timing of seedling emergence all play a significant role in determining seedling regeneration success in the face of herbivore attack. I also outline how current plant defence theories can be related to recent field observations concerning seedling acceptability and removal. In addition, I suggest that those seedling characteristics which provide the emerging plant with some degree of resistance against herbivory should be viewed as one component of a suite of plant life history traits affecting seedling regeneration success or failure. In view of the increasing importance of comparative methods in plant ecology, I propose that future research should integrate seedling acceptability with other plant life history traits thought to affect seedling regeneration. Recent work suggests that seed size may be paramount in determining regeneration success. I explore the possibility that seed size and seedling acceptability may be linked and discuss how these two factors may interact along successional gradients in plant communities.     

Correlational selection on life history traits in the pitcher-plant mosquito

We examined the short-term effects of genetic architecture on the response to correlational selection in the pitcher-plant mosquito, Wyeomyia smithii. Previously, a positive phenotypic and genetic correlation between development time and propensity to diapause had been identified. Correlational selection consisted of all four combinations of (1) fast and slow development time and (2) propensity to diapause and not diapause, 3 replicates per selection regime, for 1 to 3 generations. Response to selection in the direction of the correlation was rapid and generally consistent with the direction of selection while response orthogonal to the correlation was inconsistent and frequently in a direction opposite to the direction of selection. We conclude that genetic architecture can have a significant, and quite asymmetrical and non-directional, effect on the response to correlational selection.     

Causes of covariation of phenotypic traits among populations

Morphological and life-history traits often vary among populations of a species. Traits generally do not vary independently, but show patterns of covariation that can arise from genetic and environmental influences on phenotype. Covariance of traits may arise at an among-population level when genetically influenced traits diverge among populations in a correlated manner. Genetic correlations caused by pleiotropy and/or gene linkage can cause traits to evolve together, but among-population covariance can also arise among traits that are not genetically correlated. For example, “selective covariance” can arise when natural selection directly causes correlated change in a suite of traits. Similarly, mutation, migration, and drift may also sometimes cause correlated genetic changes among populations. Because covariation of traits among populations can arise by several different processes, the evolution of suites of traits must be interpreted with great caution. We discuss the sources of among-population covariance and illustrate one approach to identifying the sources' using data on floral traits of Dalechampia scandens (Euphorbiaceae).     

Colonization dynamics and life history traits of seven Daphnia pulex genotypes

Summary A series of experiments revealed significant differences in the potential ability of seven Daphnia pulex genotypes to colonize two lakes in which this species does not naturally reside. Life table experiments, in which individuals from each genotype were raised separately on water and natural phytoplankton from the two lakes, revealed several significant differences among genotypes in life history traits, including age and size at first reproduction, clutch size and offspring body size. Significant differences among genotypes were also found in mean genotype fitness and rate of population increase, although all genotypes were able to increase in absolute numbers. Significant genotypexlake interaction was found for several life history traits and mean fitness, indicating that the relative success of invading genotypes may depend on habitat characteristics. Enclosure experiments, in which all seven genotypes were introduced together into enclosures in both lakes, revealed that some genotypes increased greatly while others declined in relative abundance, and that the most successful genotypes differed between lakes. In addition, the most successful genotypes in the enclosures were not necessarily the genotypes that displayed the highest fitness in the life table experiments, possibly because individuals in the enclosures competed for food resources, leading to exclusion of certain genotypes.     

Generation specific life history traits of winged shape Sitobion avenae

When reared under constant conditions from egg hatch the proportion of offspring of Sitobion avenae (F.) (Aphididae: Macrosiphini) that developed into winged individuals declined from the second to fourth generation. The winged individuals produced in these three generations showed a progressive decrease in their wet and dry weights and an increase in the length of their largest embryo with generation. The trend in winged individual production and also the trends in their size and reproductive strategy appear to be programmed and are similar to the generation specific strategies that have been described in other species of aphid.     

A review of the life history,reproduction and phenology of Gracilaria

The basic life history of the red alga Gracilaria is of the three-phase Polysiphonia type but a number of species show deviations. Plants can bear both gametangia and tetrasporangia, either on separate parts of the thallus or on the same. Explanations include the in situ germination of tetraspores (allowing gametophytic thalli to be epiphytic on tetrasporophytes), the coalescence of spores or developing discs (resulting in chimaeras), mitotic recombination during cell division in the mature diploid thallus (resulting in patches of diploid male and female cells on the tetrasporophyte), a mutation eliminating the repression of female expression allowing haploid male plants to be bisexual and initial failure of cell walls to form during the development of tetraspores. Polyploids can be produced from plants with diploid gametangia. The sexes and phases are usually morphologically identical but gametophytes or their parts may be smaller. The growth rates of the sexes may differ and diploid juveniles may survive better than haploid. Neither polyploidy nor hybridization results in superior growth. The sex ratio is probably 1:1 but females may appear to be more abundant. Diploid and haploid phases are usually either about equal or diploids predominate, often depending on the type of substratum. At high latitudes reproduction peaks in late summer whereas in the tropics it may be high all year. In temperate regions growth rate is fastest and biomass highest in late summer; in the tropics peak biomass is mainly in the winter. Spermatia are effective for only a few h. Spores vary in size around 25 m, diploid ones usually being larger. Cystocarps or tetrasporangia in the field may not currently be releasing spores. In the laboratory spore release shows a diurnal rhythm, peaking during the night or day according to the species. All the above attributes are potentially important in planning and executing Gracilaria cultivation.