Experimental evolution reveals trade-offs between mating and immunity

Immune system maintenance and upregulation is costly. Sexual selection intensity, which increases male investment into reproductive traits, is expected to create trade-offs with immune function. We assayed phenoloxidase (PO) and lytic activity of individuals from populations of the Indian meal moth, Plodia interpunctella, which had been evolving under different intensities of sexual selection. We found significant divergence among populations, with males from female-biased populations having lower PO activity than males from balanced sex ratio or male-biased populations. There was no divergence in anti-bacterial lytic activity. Our data suggest that it is the increased male mating demands in female-biased populations that trades-off against immunity, and not the increased investment in sperm transfer per mating that characterizes male-biased populations.     

Behavioural differences between individuals and two populations of stickleback (Gasterosteus aculeatus)

Behavioural syndromes are correlations between behaviours in different functional contexts. Behavioural syndromes are attracting the attention of evolutionary biologists because they mean that different behaviours might not be free to evolve independently of one another. In a landmark study, Huntingford (1976) showed that individual stickleback which were bold toward predators were also aggressive toward conspecifics and active in an unfamiliar environment. Here, I revisited the activity-aggression-boldness syndrome in stickleback and tested the hypothesis that correlations between behaviours might act as evolutionary constraints. I measured a suite of behaviours on wild-caught individuals and their offspring from two different populations and calculated heritabilities and genetic correlations between the different behaviours. I found that these behaviours were phenotypically and genetically correlated in one population but not another. On average, boldness and aggression were negatively related to each other across the populations. These results suggest that behavioural syndromes don't always act as evolutionary constraints.     

Biomechanical trade-offs bias rates of evolution in the feeding apparatus of fishes

Morphological diversification does not proceed evenly across the organism. Some body parts tend to evolve at higher rates than others, and these rate biases are often attributed to sexual and natural selection or to genetic constraints. We hypothesized that variation in the rates of morphological evolution among body parts could also be related to the performance consequences of the functional systems that make up the body. Specifically, we tested the widely held expectation that the rate of evolution for a trait is negatively correlated with the strength of biomechanical trade-offs to which it is exposed. We quantified the magnitude of trade-offs acting on the morphological components of three feeding-related functional systems in four radiations of teleost fishes. After accounting for differences in the rates of morphological evolution between radiations, we found that traits that contribute more to performance trade-offs tend to evolve more rapidly, contrary to the prediction. While ecological and genetic factors are known to have strong effects on rates of phenotypic evolution, this study highlights the role of the biomechanical architecture of functional systems in biasing the rates and direction of trait evolution.     

The influence of small-scale variation in habitat on winter trout distribution and diet in an afforested catchment

(1) Habitat influences on trout distribution and diet at a reach scale were analysed over a single winter in a conifer-afforested catchment in Ireland, in an area subject to very low levels of atmospheric pollution. (2) A total of 234 salmonids were sampled from which 1194 individual prey items were recovered. Salmonids were sampled at eleven sites over a 2 km stretch of the river Douglas, a third order stream of the river Araglin in the Munster Blackwater, Co. Cork, Ireland. (3) In this winter study, water depth and flow primarily influenced trout distribution. (4) Stepwise multiple regression was used to identify the most influential environmental factors on trout metrics. Trout biomass was positively correlated to maximum depth and the percentage of pool area within sites, but density and condition were not significantly influenced by habitat variation at the reach scale. (5) The results from diet analysis suggest that trout are heterogeneous generalist feeders in winter, preying mainly on Trichoptera and Ephemeroptera species. Prey diversity in the diet of individuals throughout the study was positively correlated to the percentage of deciduous vegetation, undercut banks and coniferous vegetation, whilst the prey sub-population diversity at site scale was not related to any of the habitat variables measured.     

Habitat use and movements of plains zebra (Equus burchelli) in response to predation danger from lions

Prey species must adapt their behavior to avoid predation. Asa key prey item for lions (Panthera leo), plains zebras (Equusburchelli) were expected to respond to immediate threats posedby lions in their area. In addition, zebras were predicted toexhibit behavior tuned to reduce the potential for encounterswith lions, by modifying their movement patterns in the timesof day and habitats of greatest lion danger. We studied a populationof approximately 600 plains zebra living in Ol Pejeta Conservancy,Kenya. We found that zebra abundance on or near a grasslandpatch was lower if lions had also been observed on that patchduring the same day. Predation danger was highest in grasslandhabitat during the night, when lions were more active. Zebrasightings and global positioning system radio collar data indicatedthat zebras also reduced their use of grassland at night, insteadusing more woodland habitat. Zebras moved faster and took sharperturns in grassland at night. It is hypothesized that these moreerratic movements assist zebras in avoiding detection or captureby lions.     

Behavioural models of population growth rates: implications for conservation and prediction

Conservation biologists often wish to predict how vertebrate populations will respond to local or global changes in conditions such as those resulting from sea-level rise, deforestation, exploitation, genetically modified crops, global warming, human disturbance or from conservation activities. Predicting the consequences of such changes almost always requires understanding the population growth rate and the density dependence. Traditional means of directly measuring density dependence are often extremely difficult and have the problem that if the environment changes then it is necessary to remeasure the density dependence. We describe an alternative approach that does not require such long datasets and can be used to predict the density dependence under novel conditions. Game theory can be used to describe behavioural decisions that individuals make in response to interference, prey depletion, territorial behaviour or social dominance, and the resultant fitness consequences. It is then possible to predict how survival or reproductive output changes with population size. From this we can then make predictions about the responses of populations to environmental changes. We will illustrate how this can be applied to a range of species and a range of applied problems.     

Shifts in abundance and growth of slimy sculpin in response to changes in the predator population in an arctic Alaskan lake

Lake trout (Salvelinus namaycush) in Toolik Lake are tightly coupled to the benthos, since they have no pelagic forage fishes. Slimy sculpins (Cottus cognatus) are a prey of lake trout and the soft sediment chironomids are an important prey for the sculpin. Our previous work showed that the median size of lake trout in Toolik Lake had decreased significantly between 1977 and 1986, and smaller lake trout are likely to be less effective as sculpin predators. Using our historic data on the slimy sculpin population from 1978, we took advantage of the recent change in the predator community to examine for subsequent changes in the sculpin community. Between 1978 and 1987, the percentage of slimy sculpin caught in the soft sediments has increased (25% to 39.5%). In 1987 there was a significant difference in the mean weight of sculpin caught on different substrates. The mean individual weight of sculpins increased from the nearshore rock area to the rock/soft-sediment interface to the soft sediments. There was no difference in mean individual weight with habitat in 1978. The mean total length at age for slimy sculpins during this time has also increased significantly. We suggest that the risk of predation while foraging in the soft sediments has declined. The increased use of the soft sediment area appears to have contributed to their increased growth, likely due to greater food abundance.     

Modelling trade-offs between mortality and fitness associated with persistent blood feeding by mosquitoes

Feeding persistence of mosquitoes (that tendency to attack again within a short period if a first meal is incomplete) is a key determinant of feeding success (defined here as blood-meal volume), especially with regard to mosquitoes that attack defensive hosts. Mortality risk from the vertebrate host is one cost that may be expected to create a trade-off against increased fecundity from second and subsequent meals depending on the availability of blood sources and on the probability of survival until the next gonotrophic cycle. We initially used stochastic simulation models to examine potential trade-offs in fecundity and survival for persistent and nonpersistent lines of mosquitoes that acquire blood from potentially dangerous hosts. In these models, a given strategy, i.e. persistence or the lack thereof, was fixed for the lifetime of the mosquito. Persistent blood feeding paid off relative to nonpersistent mosquitoes when feeding-associated mortality was relatively low to moderate and when the probability of obtaining blood on a given attempt was relatively low. On the other hand, the nonpersistent strategy tended to give relatively higher fecundity when the chances of obtaining blood were high. We also examined the implications of persistence as a feeding strategy for mosquitoes that could exhibit this behaviour in a flexible manner over their lifetime in response to state. We solved for optimal feeding behaviours with a stochastic dynamic programming model set up along similar lines to the simulation model. Results from this approach lend support to the idea that the threshold blood-meal size at which mosquitoes should continue to feed should change relative to the age of the mosquito. The modelling approaches used in this study identified important areas of future study for mosquito blood-feeding behaviour.     

Habitat use and life history of brown trout (Salmo trutta) and Arctic charr (Salvelinus alpinus) in some low acidity lakes in central Norway

Habitat utilization and the life history of browntrout Salmo trutta and Arctic charr Salvelinus alpinus were investigated in fivesympatric populations and five allopatric brown troutpopulations in Høylandet catchment, a atmosphaericlow deposition area in Mid Norway. There was asignificant inverse correlation in abundance ofepibenthic Arctic charr and brown trout in theselakes, indicating that the latter species is dominant.The largest numbers of sympatric brown trout andArctic charr were caught in epibenthic habitat. In twolakes, brown trout to some extent also occurredpelagically, while pelagic individuals of Arctic charrwere found in all five lakes. The main food items forboth epibenthic and pelagic brown trout wereterrestrial surface insects and chironomid pupae.Zooplankton was the primary food item for Arctic charrin both habitats. Although the age distribution wasvery different in the populations, neither speciesseem to suffer from recruitment failure. There was nosignificant difference in survival rates betweensympatric populations of brown trout and Arctic charr.We found a significant inverse correlation betweenepibenthic catches of brown trout and the mean weightof 4+ fish, the most abundant age group. However, ifusing weight data for three-year-old fish, no suchrelationship was found for Arctic charr. Brown troutand Arctic charr reached asymptotic lengths of197–364 mm and 259–321 mm, respectively. Both speciestypically reached sexual maturity at age 2–3, and nomaturation-induced mortality was evident. We concludethat fish populations in Høylandet lakes areregulated throughout their lifes by inter- andintraspecific competition.     

Mating opportunity and the evolution of sex-specific mortality rates in a butterfly

Life history theory predicts that organisms should only invest resources into intrinsic components of life span to the degree that it pays off in terms of reproductive success. Here, we investigate if the temporal distribution of mating opportunities may have influenced the evolution of intrinsic mortality rates in the butterfly Pararge aegeria (Satyrinae). In this species, females mate only once and the frequency of male mating opportunities depends on the temporal emergence pattern of virgin females. As expected, in a population from Madeira where females emerge continuously throughout the year, there was no sex difference in adult life span, while in a Swedish population with synchronised female emergence, males had significantly shorter life spans compared to females. A logistic mortality model provided the best fit to the observed change in age-specific mortality and all categories reached an asymptotic mortality rate of a similar magnitude. However, the Swedish males reached this mortality plateau more rapidly than the other categories. External mortality, due to water and food limitation, affected the pattern of sex-specific mortality but males from Sweden still had higher rates of mortality compared to all other categories. We argue that selection on male longevity is likely to be weaker in Sweden because under synchronised emergence, all females emerge and mate within a short period of time, after which male reproductive value will quickly approach zero. On Madeira, however, male reproductive value decrease more slowly with age since the probability of finding a receptive female is constant over the year. Received: 29 July 1999 / Accepted: 23 August 1999     

Positive correlations between molecular and morphological rates of evolution

The existence of positive associations between rates of molecular and morphological evolution (calculated from branch lengths of phylogenetic trees reconstructed using molecular and morphological characters, respectively) is important to issues of neutrality in sequence evolution, phylogenetic reconstructions assuming neutrality, and evolutionary genotype-phenotype mapping. Rates correlate positively when including branches leading to extant species (tips). Excluding tips, trends are similar, but statistical significances decrease systematically. This is due to (a) lower statistical power (excluding tips reduces sample sizes), and (b) rates are solely calculated from inaccurately reconstructed character states of extinct ancestral species, and this noise decreases correlation strengths. Correlations between molecular and morphological rates of evolution increase as more morphological characters are included for phylogenetic reconstruction. Sequence lengths apparently affect correlations along similar principles. Analyses of plant phylogenies confirm those from animals: sampling biases decrease correlations between molecular and morphological rates of evolution. Results confirm that genotype and phenotype are linked, and suggest adaptive components for molecular evolution. The discussion stresses the difficulties associated with analyses and conclusions based on data deduced from phylogenetic reconstruction.     

Sibling competition and the evolution of growth rates in birds

Variation among bird species in growth rates is traditionally attributed to differences in energy availability and developmental mode. However, the extent and form of competition among siblings for limited food resources may also be an important determinant. Kin-selection-based models of intrabrood competition suggest that nestling growth rates should be highest in those species in which siblings are likely to be less genetically related to one another (half-sibs rather than full-sibs). We test this novel prediction using the frequency of multiple paternity as an index of average sib relatedness within broods. As predicted, we find a significant positive association between the rate of multiple paternity within broods and nestling growth rates. Furthermore, this holds true when we control for the effects of variation in other factors that may be associated with variation in growth rate, such as body size, brood size, mating system and the form of parental care. We suggest, therefore, that variation in growth rate among bird species is not simply dependent on proximate ecological and developmental factors but is also strongly influenced by interactions, over an evolutionary time-scale, among kin.