Variation in life-history characteristics among populations of North American wood turtles: a view from the north

Life-history traits such as age at maturity, body size and clutch size tend to vary across a species' distribution. The purpose of our study was to describe the demography of a newly discovered population of North American wood turtles Glyptemys insculpta at the species' northern range limit, and to compare our findings to those of other studies to test hypotheses about adaptive life-history variation. Turtles were hand-captured from May to October 2005 and 2006 along a 4.5 km stretch of river located in the Sudbury District, ON, Canada (46°N). Fifty-five captured individuals provided a population density estimate of 1.3 turtles/100 m of river. Juveniles comprised 35% of wood turtles captured, and growth ring counts (i.e. age estimates) indicated recruitment in each of the past 11 years. Among populations, we found a nonlinear pattern in body size variation with the largest turtles in the north, smallest turtles in the centre of the range, and intermediate-sized turtles in the south. This nonlinear pattern in body size was reflected in clutch size variation. Selective pressures to overcome years of low recruitment may have resulted in larger body sizes and hence large clutch sizes at northern latitudes while conspecifics at southern latitudes can achieve larger body sizes because they live in a more productive environment. Population density decreased with latitude, likely as a result of a gradient in habitat productivity.     

The timing of life-history events in a changing climate

Although empirical and theoretical studies suggest that climate influences the timing of life-history events in animals and plants, correlations between climate and the timing of events such as egg-laying, migration or flowering do not reveal the mechanisms by which natural selection operates on life-history events. We present a general autoregressive model of the timing of life-history events in relation to variation in global climate that, like autoregressive models of population dynamics, allows for a more mechanistic understanding of the roles of climate, resources and competition. We applied the model to data on 50 years of annual dates of first flowering by three species of plants in 26 populations covering 4 degrees of latitude in Norway. In agreement with earlier studies, plants in most populations and all three species bloomed earlier following warmer winters. Moreover, our model revealed that earlier blooming reflected increasing influences of resources and density-dependent population limitation under climatic warming. The insights available from the application of this model to phenological data in other taxa will contribute to our understanding of the roles of endogenous versus exogenous processes in the evolution of the timing of life-history events in a changing climate.     

The physiology of life-history trade-offs: experimental analysis of a hormonally induced life-history trade-off in Gryllus assimilis

Abstract Adult Gryllus assimilis given an analog of juvenile hormone exhibited reduced flight muscles and enlarged ovaries similar to those found in naturally occurring flightless individuals of species that are polymorphic for dispersal capability. Control and hormone-treated (flightless) G. assimilis did not differ in the amount of food consumed or assimilated on any of three diets that differed in nutrient quantity. Thus, enhanced ovarian growth of flightless individuals resulted from increased allocation of internal nutrients to reproduction (i.e., a trade-off) rather than from increased acquisition of nutrients. Compared with flight-capable controls, flightless G. assimilis also had reduced whole-organism respiration, reduced respiration of flight muscles, and reduced lipid and triglyceride (flight fuel) reserves. These differences are remarkably similar to those between naturally occurring flightless and flight-capable morphs of other Gryllus species. Results collectively suggest that the increased allocation of nutrients to ovarian growth in flightless G. assimilis and other Gryllus species results from reduced energetic costs of flight muscle maintenance and/or the biosynthesis or acquisition of lipids. Reduction in these energetic costs appears to be an important driving force in the evolution of flightlessness in insects. Respiratory metabolism associated with flight capability utilizes an increasing proportion of the energy budget of crickets as the quantity of nutrients in the diet is decreased. This leads to a magnification of greater ovarian growth of flightless versus flight-capable individuals on nutrient-poor diets.     

The ten Hox genes of the millipede Glomeris marginata

We have isolated the ten Hox genes from the pill millipede Glomeris marginata (Myriapoda:Diplopoda). All ten genes are expressed in characteristic Hox-gene-like expression patterns. The register of Hox gene expression borders is conserved and the expression profiles show that the anterior-most limb-bearing segment in arthropods (antennal/cheliceral segment) does not express any Hox gene, while the next segment (intercalary/second-antennal/premandibular/pedipalpal segment) does express Hox genes. The Hox expression patterns in this millipede thus support the conclusion that all arthropods possess a deuterocerebral segment. We find that there is an apparent posterior shift of Hox gene expression domains dorsally relative to their ventral patterns, indicating that the decoupling of dorsal and ventral segmentation is not restricted to the level of segment polarity genes but apparently includes the Hox genes. Although the mechanism for the decoupling of dorsal and ventral segmentation remains unsolved, the decoupling must be at a level higher in the hierarchy than that of the segment polarity and Hox genes. The expression patterns of Ultrabithorax and abdominal-A suggest a correlation between the function of these genes and the delayed outgrowth of posterior trunk appendages. This delay may be caused by an assumed repressor function of Ultrabithorax, which might partially repress the activation of the Distal-less gene. The Glomeris fushi tarazu gene is expressed in a Hox-like domain and in the developing central nervous system, but not in segmental stripes such as has been reported in another myriapod species, the centipede Lithobius. In contrast to the Lithobius fushi tarazu gene, there is no indication for a role in segment formation for the millipede fushi tarazu gene, suggesting that fushi tarazu first acquired its segmentation function in the lineage of the insects.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Dispersal-related life-history trade-offs in a butterfly metapopulation

1. Recent studies on butterflies have documented apparent evolutionary changes in dispersal rate in response to climate change and habitat change. These studies often assume a trade-off between dispersal rate (or flight capacity) and reproduction, which is the rule in wing-dimorphic species but might not occur equally in wing-monomorphic species such as butterflies. 2. To investigate the relationship between dispersal rate and fecundity in the Glanville fritillary butterfly Melitaea cinxia we recorded lifetime individual movements, matings, ovipositions, and maximal life span in a large (32 x 26 m) population cage in the field. Experimental material was obtained from 20 newly established and 20 old local populations within a large metapopulation in the Aland Islands in Finland. 3. Females of the Glanville fritillary from newly established populations are known to be more dispersive in the field, and in the cage they showed significantly greater mobility, mated earlier, and laid more egg clutches than females from old populations. The dispersive females from new populations exhibited no reduced lifetime fecundity in the cage, but they had a shorter maximal life span than old-population females. 4. These results challenge the dispersal-fecundity trade-off for nonmigratory butterflies but instead suggest a physiological trade-off between high metabolic performance and reduced maximal life span. High metabolic performance may explain high rates of dispersal and oviposition in early life. 5. In fragmented landscapes, an ecological trade-off exists between being more dispersive and hence spending more time in the landscape matrix vs. having more time for reproduction in the habitat. We estimate with a dispersal model parameterized for the Glanville fritillary that the lifetime egg production is 4% smaller on average in the more dispersive butterflies in a representative landscape, with much variation depending on landscape structure in the neighbourhood of the natal patch, from--26 to 45% in the landscape analysed in this paper.     

The life-history basis of behavioural innovations

The evolutionary origin of innovativeness remains puzzling because innovating means responding to novel or unusual problems and hence is unlikely to be selected by itself. A plausible alternative is considering innovativeness as a co-opted product of traits that have evolved for other functions yet together predispose individuals to solve problems by adopting novel behaviours. However, this raises the question of why these adaptations should evolve together in an animal. Here, we develop the argument that the adaptations enabling animals to innovate evolve together because they are jointly part of a life-history strategy for coping with environmental changes. In support of this claim, we present comparative evidence showing that in birds, (i) innovative propensity is linked to life histories that prioritize future over current reproduction, (ii) the link is in part explained by differences in brain size, and (iii) innovative propensity and life-history traits may evolve together in generalist species that frequently expose themselves to novel or unusual conditions. Combined with previous evidence, these findings suggest that innovativeness is not a specialized adaptation but more likely part of a broader general adaptive system to cope with changes in the environment.     

The millipedes of a Cheshire wood

The density and surface activity of the millipedes in a quarter of an acre of a sycamore ash wood are described. Seven species were extracted by Tullgren funnels from samples of soil and litter over five years and were also caught in pitfall traps during a further two years; four other species occurred occasionally in the traps. Each square metre of the site supported 100 individuals over the winter, rising to 300 in the summer. Of these, 85 % belonged to three species, Iulus scandinavius, Polydesmus angustus and P. denticulatus .
Male I. scandinavius become adult in either the ninth, tenth or eleventh stadium; females in the tenth and eleventh. Eggs are laid in spring and these take three years to become adults which breed and then die. The majority of Polydesmus spp. in the samples are young belonging to the first six of the eight free-living stadia. The adults fell into the traps in the summer and newly emerged young appear in the samples at this time. They overwinter in their first year mainly as fifth stadia; some might reach maturity (eighth stadia) in the summer following, but it is not certain that they could breed at this time.
The pattern of dispersion of lulus is fairly even and is correlated with the distribution of leaf litter but the Polydesmus spp. are highly aggregated. All stadia of lulus fall into the traps but only the last two of Polydesmus . The aggregation of Polydesmus spp. appears to be correlated with the relative inactivity of the younger stadia. The estimates of density of Polydesmus spp. are unreliable because of their aggregation but those of lulus have determinable limits and it is possible to derive rough though meaningful standing crop, production and life-table data. The overwintering standing crop of lulus consists of the survivors of three generations of 5, 2.2 and 1.4% of the original eggs laid; it has a fresh weight of about 1.25 g and a production in the order of 1.5–2.5 g/m².     

The evolution of population stability as a by-product of life-history evolution

Proposed mechanisms for the evolution of population stability include group selection through longterm persistence, individual selection acting directly on stability determining the demographic parameters, and the evolution of stability as a by-product of life-history evolution. None of these hypotheses currently has clear empirical support. Using two sets of Drosophila melanogaster populations, we provide experimental evidence of stability evolving as a correlated response to selection on traits not directly related to demography. Four populations (FEJs) were selected for faster development and early reproduction for 125 generations, and the other four (JBs) were ancestral controls. All FEJ and JB populations have been maintained on discrete generations at moderate density, thus eliminating differential selection on stability determining demographic parameters. We derived eight small populations from each FEJ and JB population, and subjected four small populations each to either stabilizing or destabilizing food regimes. Census data on these 64 small populations over 20 generations clearly showed that the FEJ populations have significantly less temporal fluctuations in their numbers in both food regimes compared to their controls. This greater stability of the FEJ populations is probably a by-product of the evolution of reduced fecundity and pre-adult survivorship, as a correlated response to selection for rapid development.     

Identification of a fluorescent compound in the cuticle of the train millipede Parafontaria laminata armigera

The train millipede (Parafontaria laminata armigera) emits a blue fluorescence (λ(max)=455 nm) under black light (350 nm). The isolated fluorescent compound from the cuticle of P. laminata armigera was identified as pterin-6-carboxylic acid. The structure of this compound was identified by fluorescent, HPLC, and mass spectrometric (ESI-ion trap MS) analyses, and then compared with an authentic sample.     

Inbreeding depression along a life-history continuum in the great tit

Inbreeding resulting from the mating of two related individuals can reduce the fitness of their progeny. However, quantifying inbreeding depression in wild populations is challenging, requiring large sample sizes, detailed knowledge of life histories and study over many generations. Here we report analyses of the effects of close inbreeding, based on observations of mating between relatives, in a large, free-living noninsular great tit (Parus major) population monitored over 41 years. Although mating between close relatives (f > or = 0.125) was rare (1.0-2.6% of matings, depending on data set restrictiveness), we found pronounced inbreeding depression, which translated into reduced hatching success, fledging success, recruitment to the breeding population and production of grand offspring. An inbred mating at f = 0.25 had a 39% reduction in fitness relative to that of an outbred nest, when calculated in terms of recruitment success, and a 55% reduction in the number of fledged grand offspring. Our data show that inbreeding depression acts independently at each life-history stage in this population, and hence suggest that estimates of the fitness costs of inbreeding must focus on the entire life cycle.     

Spermatogeny as a life-history index in parasitoid wasps

Reproduction is a major life-history trait but it has been studied mostly in relation to female reproductive effort. Recently, an ovigeny index using the proportion of oocytes ready to be oviposited at eclosion has been proposed for female insect parasitoids. Here, we propose a spermatogeny index for male parasitoid wasps. Prospermatogenic species have an index of 1, have all their spermatozoids mature at emergence and do not produce more later in life. At the other end of the spectrum, synspermatogenic species have no spermatozoids at emergence and produce them later in life. The level of spermatogeny should be linked to several other life-history parameters such as longevity, size, nutrition, distribution of mating opportunities and dispersion before and after mating. Data presented for some parasitoid species support the presence of variability in this male life-history parameter.     

Predator induced life-history shifts in a freshwater cladoceran

Summary Life-history theory predicts that maturity and resource allocation patterns are highly sensitive to selective predation. Under reduced adult survival, selection will favour genotypes capable of reproducing earlier, at a smaller size and with a higher reproductive effort. When exposed to water that previously held fish, (size selective predators which prefer larger Daphnia), individuals of Daphnia hyalina reproduced earlier, at a smaller size and had a higher reproductive investment. Hence the prey was able to switch its life history pattern in order to become less susceptible to predation by a specific predator. The cue that evokes the prey response is a chemical released by the predator.     

The nature and dynamics of world religions: a life-history approach

In contrast with tribal and archaic religions, world religions are characterized by a unique emphasis on extended prosociality, restricted sociosexuality, delayed gratification and the belief that these specific behaviours are sanctioned by some kind of supernatural justice. Here, we draw on recent advances in life history theory to explain this pattern of seemingly unrelated features. Life history theory examines how organisms adaptively allocate resources in the face of trade-offs between different life-goals (e.g. growth versus reproduction, exploitation versus exploration). In particular, recent studies have shown that individuals, including humans, adjust their life strategy to the environment through phenotypic plasticity: in a harsh environment, organisms tend to adopt a ‘fast'' strategy, pursuing smaller but more certain benefits, while in more affluent environments, organisms tend to develop a ‘slow'' strategy, aiming for larger but less certain benefits. Reviewing a range of recent research, we show that world religions are associated with a form of ‘slow'' strategy. This framework explains both the promotion of ‘slow'' behaviours such as altruism, self-regulation and monogamy in modern world religions, and the condemnation of ‘fast'' behaviours such as selfishness, conspicuous sexuality and materialism. This ecological approach also explains the diffusion pattern of world religions: why they emerged late in human history (500–300 BCE), why they are currently in decline in the most affluent societies and why they persist in some places despite this overall decline.     

Ecological immunogenetics of life-history traits in a model amphibian

Major histocompatibility complex (MHC) genes determine immune repertoires and social preferences of vertebrates. Immunological regulation of microbial assemblages associated with individuals influences their sociality, and should also affect their life-history traits. We exposed Xenopus laevis tadpoles to water conditioned by adult conspecifics. Then, we analysed tadpole growth, development and survivorship as a function of MHC class I and class II peptide-binding region amino acid sequence similarities between tadpoles and frogs that conditioned the water to which they were exposed. Tadpoles approached metamorphosis earlier and suffered greater mortality when exposed to immunogenetically dissimilar frogs. The results suggest that developmental regulatory cues, microbial assemblages or both are specific to MHC genotypes. Tadpoles may associate with conspecifics with which they share microbiota to which their genotypes are well adapted.     

Pathogen-induced rapid evolution in a vertebrate life-history trait

Anthropogenic factors, including climate warming, are increasing the incidence and prevalence of infectious diseases worldwide. Infectious diseases caused by pathogenic parasites can have severe impacts on host survival, thereby altering the selection regime and inducing evolutionary responses in their hosts. Knowledge about such evolutionary consequences in natural populations is critical to mitigate potential ecological and economic effects. However, studies on pathogen-induced trait changes are scarce and the pace of evolutionary change is largely unknown, particularly in vertebrates. Here, we use a time series from long-term monitoring of perch to estimate temporal trends in the maturation schedule before and after a severe pathogen outbreak. We show that the disease induced a phenotypic change from a previously increasing to a decreasing size at maturation, the most important life-history transition in animals. Evolutionary rates imposed by the pathogen were high and comparable to those reported for populations exposed to intense human harvesting. Pathogens thus represent highly potent drivers of adaptive phenotypic evolution in vertebrates.