The evolution of reproductive strategies in dasyurid marsupials: implications of molecular phylogeny

Dasyurid marsupials show a remarkable diversity of reproductive patterns ranging from aseasonal polyoestry to restricted annual breeding in which males synchronously die after a brief mating season. Previous studies have categorized dasyurid reproduction into six strategies, defined on the basis of five life-history characters. We provide an up-to-date summary of reproductive traits in dasyurid species and examine the evolution of these characters on a phylogeny for the family recently obtained from DNA sequence data. Our results suggest that reproductive evolution in modern dasyurids is characterized by a basal separation of subfamily lineages employing Strategy II (monoestrous females, restricted breeding season, 11 months to maturity; Dasyurinae) and Strategy V (polyoestrous females, extended breeding season, 8–11 months to maturity; Sminthopsinae). Strategies I (male die-off) and III (facultative polyoestry) appear to have arisen several times from Strategy II or V ancestors, and Strategy IV appears to have arisen within Sminthopsis from a Strategy V ancestor. Strategy VI (aseasonal breeding) has arisen independently in each of the four major dasyurid lineages (tribes), and is highly (but not perfectly) correlated with New Guinean endemism. This scenario is not strongly affected if reproductive characters are optimized on an alternative phylogeny more consistent with morphology-based opinions on species relationships. When evaluated in light of current habitat associations and geographic distributions, the reproductive data suggest that the Miocene diversification of modern dasyurids may have been correlated with the invasion of dry forest or woodland habitats.     

Variation in fecundity and other reproductive traits in freshwater mussels

1. Life histories of the highly diverse and endangered North American freshwater mussel fauna are poorly known. We investigated reproductive traits of eight riverine mussel species in Alabama and Mississippi, U.S.A.: Amblema plicata, Elliptio arca, Fusconaia cerina, Lampsilis ornata, Obliquaria reflexa, Pleurobema decisum, Quadrula asperata and Q. pustulosa, and compare our results with existing life history information for other species. 2. These eight species had reproductive traits characteristic of large, outcrossing populations: hermaphrodites were rare, we found no evidence of protandry, and sex ratios were even or slightly male‐biased. 3. Age at sexual maturity varied among species, ranging from <1 to 2 years for L. ornata to 3–9 years for Q. asperata. In all species, most mature females participated in reproduction and fertilisation success was high. 5. Fecundity was related positively to both length and age, but length was the best predictor. In six species, fecundity increased exponentially with increasing size; in two species the rate of increase in fecundity declined in larger animals. In four species, fecundity declined in older animals. These latter results indicate weak reproductive senescence; however, in all species, older individuals continued to produce large numbers of offspring. Mean annual fecundity differed widely among species ranging from 9647 to 325 709. Within‐species differences in fecundity were found among rivers and among populations within a river. 6. The wide variation in reproductive traits among species indicates the existence of widely divergent life history strategies in freshwater mussels.     

Comparing and contrasting life history variation in four aphid hyperparasitoids

1. In primary parasitoids, significant differences in life history and reproductive traits are observed among parasitoids attacking different stages of the same host species. Much less is known about hyperparasitoids, which attack different stages of primary parasitoids. 2. Parasitoids exploit hosts in two different ways. Koinobionts attack hosts that continue feeding and growing during parasitism, whereas idiobionts paralyse hosts before oviposition or attack non‐growing host stages, e.g. eggs or pupae. 3. Koino‐/idiobiosis in primary parasitoids are often associated with different expression of life history trade‐offs, e.g. endo‐ versus ectoparasitism, high versus low fecundity and short versus long life span. 4. In the present study, life history parameters of two koinobiont endoparasitic species (Alloxysta victrix; Syrphophagus aphidivorus), and two idiobiont ectoparasitic species (Asaphes suspensus; Dendrocerus carpenteri) of aphid hyperparasitoids were compared. These hyperparasitoids attack either the parasitoid larva in the aphid before it is killed and mummified by the primary parasitoid or the parasitoid prepupa or pupa in the dead aphid mummy. 5. There was considerable variation in reproductive success and longevity in the four species. The idiobiont A. suspensus produced the most progeny by far and had the longest lifespan. In contrast, the koinobiont A. victrix had the lowest fecundity. Other developments and life history parameters in the different species were variable. 6. The present results reveal that there was significant overlap in life history and reproductive traits among hyperparasitoid koinobionts and idiobionts, even when attacking the same host species, suggesting that selection for expression of these traits is largely association specific.     

Optimal time to patency in parasitic nematodes: host mortality matters

We develop an optimality model based on classical epidemiological models to investigate the optimal time to patency in parasitic nematodes in relation to host mortality and parasite mortality. We found that the optimal time to patency depends on both host longevity and prepatent mortality of nematodes. We tested our models using a comparative analysis of the relationships between nematode time to patency, nematode mortality and host mortality. Although we confirmed the importance of prepatent mortality, we also found a significant positive influence of host mortality. Host mortality rate affects parasite survivorship and life history strategies in the same way that habitat-specific mortality regimes drive the evolution of life histories in free-living organisms.     

Fecundity of the American eel Anguilla rostrata at 45° N in Maine, U.S.A.

The northern portion of the geographic range of the American eel Anguilla rostrata may contribute a great proportion of the reproductive potential to this panmictic species because of apparent increases in average female size and female percentage with latitude. The regressions of fecundity on body length and on body weight of 63 female eels captured at about 45° N latitude on their spawning migration to the sea were log F= 1·2601 + 2·9642 log L and log F= 4·1646+0·9153 log W , where F is fecundity, L is total length (cm), and W is total weight (g). Length and weight each explained about 90% of the variation in fecundity. Estimates of fecundity from counts of aliquots of eggs ranged from 1·84 million to 19·92 million eggs for eels ranging in length from 45 to 113 cm, nearly the range of sizes of migrating females reported in the literature. Fecundities of the American eel were greater than reported in one study at about 37° N and greater than reported for the European eel, A. anguilla , shortfin eel, A. australis , and longfin eel, A. dieffenbachii . If a geographic cline in fecundity does exist in American eels, it is established anew each generation because the species forms a single panmictic population.     

Egg maturation and oviposition strategy of a capital breeder,Cleorodes lichenaria,feeding on lichens at the larval stage

Abstract 1. Capital breeders rely solely on resources acquired before breeding, whereas income breeders may obtain considerable amounts of resources following the commencement of reproduction. In income breeders oviposition occurs over a longer time period with a small number of eggs ready to be laid at the start of adult life, whereas capital breeders reproduce more rapidly and contain higher numbers of mature eggs at eclosion relative to potential fecundity. 2. This study explored egg maturation and oviposition strategy of a nocturnal geometrid moth, Cleorodes lichenaria. It was predicted, based on the long larval period of C. lichenaria and the known biology of other geometrids, that this lichen‐feeding geometrid moth concentrates egg production at the beginning of its breeding period and that external nutrients have a limited role on its fecundity. 3. Approximately 46% of eggs were ready to be laid in newly hatched females and approximately 40% of their potential fecundity was realised during the first night, after which the number and the weight of eggs decreased steadily. Feeding or drinking did not increase the fecundity of females. Pupal mass correlated positively with realised fecundity and ovigeny index. 4. Reproduction of C. lichenaria is based solely on larval‐derived resources. It is suggested that females are able to resorb thoracic tissues to increase fecundity and that this ability is probably linked to relatively long lifespan and low ovigeny index of this species, characteristics rarely observed in other geometrid moths.     

Cancer brings forward oviposition in the fly Drosophila melanogaster

Hosts often accelerate their reproductive effort in response to a parasitic infection, especially when their chances of future reproduction decrease with time from the onset of the infection. Because malignancies usually reduce survival, and hence potentially the fitness, it is expected that hosts with early cancer could have evolved to adjust their life‐history traits to maximize their immediate reproductive effort. Despite the potential importance of these plastic responses, little attention has been devoted to explore how cancers influence animal reproduction. Here, we use an experimental setup, a colony of genetically modified flies Drosophila melanogaster which develop colorectal cancer in the anterior gut, to show the role of cancer in altering life‐history traits. Specifically, we tested whether females adapt their reproductive strategy in response to harboring cancer. We found that flies with cancer reached the peak period of oviposition significantly earlier (i.e., 2 days) than healthy ones, while no difference in the length and extent of the fecundity peak was observed between the two groups of flies. Such compensatory responses to overcome the fitness‐limiting effect of cancer could explain the persistence of inherited cancer‐causing mutant alleles in the wild.     

Indiscriminate nursing in communal breeders: a role for genomic imprinting

Abstract In several communally nesting mammal species, females indiscriminately nurse each others' offspring. Previous hypotheses have suggested that the inability to recognize one's own young during lactation is the result of costs incurred from recognition errors. Here, we propose an alternative hypothesis based on sexual conflict theory and genomic imprinting. In polygynous species, males copulate with several females that may later breed communally. Under such conditions, males benefit from indiscriminate nursing of all their offspring and the reduced risk of female infanticide. This may have selected for paternally expressed genes that suppress kin recognition during lactation.     

The digestive tract and life history of small mammals

The type of food, differentiation of the large intestine and stomach, and methane production, as well as life history data, are considered in Insectivora, Rodentia and Lagomorpha. When food containing plant cell wall material is eaten, there is either a differentiation of the stomach or the large intestine. In animals with low body mass and little differentiation of the gastrointestinal tract, methane production is low, but structures essential for microbial digestion of plant cell wall material, such as haustration of the colon or formation of a caecum, can be found in many methane‐producers. Animals with a body mass < 500 g and a weaning time < 20 days are non‐producers of methane. Establishment of a balanced microbial population in the gastrointestinal tract requires some time. Many non‐producers of methane wean their young in < 10 days, but many producers need > 50 days for the weaning process. Caviomorpha, Thryonomyidae and Hystricidae seem to have ‘opened the door’ to the use of low quality food by microbial fermentation, but some of them have to ‘pay’ for this extension of the food range by an extended weaning period, which also means an extended dependency on the mother.     

Variance in male reproductive success and sexual size dimorphism in pinnipeds: testing an assumption of sexual selection theory

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Age‐specific fecundity of mammalian populations: A test of three mathematical models

Empirical age‐specific fecundity distributions are often based on small samples and hence include high levels of sampling error, particularly at the older ages. One solution to this problem is to smooth the distributions using appropriate models. The aim of this article is to compare the utility of three models for smoothing and/or graduating these distributions. The three models examined are 1) the Gamma distribution, 2) the Hadwiger function, and 3) the Brass polynomial. Test data sets consist of four types of primates (including humans), Asian elephants, and Przewalski’s horse (an extinct species). The results indicate that all three models work well with a variety of mammalian data. The simplest of these models, the Brass polynomial, cannot be rejected based on available data and appears to be the optimum choice. Zoo Biol 20:487–499, 2001. © 2002 Wiley‐Liss, Inc.     

Life histories and the evolution of cooperative breeding in mammals

While the evolution of cooperative breeding systems (where non-breeding helpers participate in rearing young produced by dominant females) has been restricted to lineages with socially monogamous mating systems where coefficients of relatedness between group members are usually high, not all monogamous lineages have produced species with cooperative breeding systems, suggesting that other factors constrain the evolution of cooperative breeding. Previous studies have suggested that life-history parameters, including longevity, may constrain the evolution of cooperative breeding. Here, we show that transitions to cooperative breeding across the mammalian phylogeny have been restricted to lineages where females produce multiple offspring per birth. We find no support for effects of longevity or of other life-history parameters. We suggest that the evolution of cooperative breeding has been restricted to monogamous lineages where helpers have the potential to increase the reproductive output of breeders.     

Allometry of human fertility and energy use

The flux of energy and materials constrains all organisms, and allometric relationships between rates of energy consumption and other biological rates are manifest at many levels of biological organization. Although human ecology is unusual in many respects, human populations also face energetic constraints. Here we present a model relating fertility rates to per capita energy consumption rates in contemporary human nations. Fertility declines as energy consumption increases with a scaling exponent of ?1/3 as predicted by allometric theory. The decline may be explained by parental trade‐offs between the number of children and the energetic investment in each child. We hypothesize that the ?1/3 exponent results from the scaling properties of the networked infrastructure that delivers energy to consumers. This allometric analysis of human fertility offers a framework for understanding the demographic transition to smaller family sizes, with implications for human population growth, resource use and sustainability.     

Life history buffering in Antarctic mammals and birds against changing patterns of climate and environmental variation

The consequences of warming for Antarctic long‐lived organisms depend on their ability to survive changing patterns of climate and environmental variation. Among birds and mammals of different Antarctic regions, including emperor penguins, snow petrels, southern fulmars, Antarctic fur seals and Weddell seals, we found strong support for selection of life history traits that reduce interannual variation in fitness. These species maximize fitness by keeping a low interannual variance in the survival of adults and in their propensity to breed annually, which are the vital rates that influence most the variability in population growth rate (λ). All these species have been able to buffer these rates against the effects of recent climate‐driven habitat changes except for Antarctic fur seals, in the Southwest Atlantic. In this region of the Southern Ocean, the rapid increase in ecosystem fluctuation, associated with increasing climate variability observed since 1990, has limited and rendered less predictable the main fur seal food supply, Antarctic krill. This has increased the fitness costs of breeding for females, causing significant short‐term changes in population structure through mortality and low breeding output. Changes occur now with a frequency higher than the mean female fur seal generation time, and therefore are likely to limit their adaptive response. Fur seals are more likely to rely on phenotypic plasticity to cope with short‐term changes in order to maximize individual fitness. With more frequent extreme climatic events driving more frequent ecosystem fluctuation, the repercussions for life histories in many Antarctic birds and mammals are likely to increase, particularly at regional scales. In species with less flexible life histories that are more constrained by fluctuation in their critical habitats, like sea‐ice, this may cause demographic changes, population compensation and changes in distribution, as already observed in penguin species living in the Antarctic Peninsula and adjacent islands.     

Fecundity and juvenile survival of shape Bembidion lampros and Tachyporus hypnorum

Fecundity and development from first instar larvae to adult beetles were studied for Bembidion lampros Herbst (Coleoptera; Carabidae) and Tachyporus hypnorum F. (Coleoptera; Staphylinidae). Both species reproduce in arable fields, where they are known to contribute to the natural control of cereal aphids. They are univoltine and have similar life cycles and development times. T. hypnorum females laid approximately 3 times more eggs than B. lampros females. Relative lifetime fecundity of B. lampros was very low with an average of 10 eggs per female. Based on the time where half of the populations had dispersed from the overwintering sites into an arable field B. lampros started egg-lay 100°D (°C>9) and T. hypnorum 200°D (°C>3) after dispersal. Mortality in the juvenile stages was found to be an important factor in the population dynamics of both B. lampros and T. hypnorum. The survival from first instar larvae to adults was on average 34% for B. lampros independent of soil type, 21% for T. hypnorum developed in sandy soil and 49% when developed in clayey soil under semi-field conditions. Enlargement of B. lampros and T. hypnorum populations to increase the natural control of aphids during their establishment in cereal fields may be possible if the conditions for juvenile development are improved.     

Fecundity selection theory: concepts and evidence

Fitness results from an optimal balance between survival, mating success and fecundity. The interactions between these three components of fitness vary depending on the selective context, from positive covariation between them, to antagonistic pleiotropic relationships when fitness increases in one reduce the fitness of others. Therefore, elucidating the routes through which selection shapes life history and phenotypic adaptations via these fitness components is of primary significance to understanding ecological and evolutionary dynamics. However, while the fitness components mediated by natural (survival) and sexual (mating success) selection have been debated extensively from most possible perspectives, fecundity selection remains considerably less studied. Here, we review the theoretical basis, evidence and implications of fecundity selection as a driver of sex‐specific adaptive evolution. Based on accumulating literature on the life‐history, phenotypic and ecological aspects of fecundity, we (i) suggest a re‐arrangement of the concepts of fecundity, whereby we coin the term ‘transient fecundity’ to refer to brood size per reproductive episode, while ‘annual’ and ‘lifetime fecundity’ should not be used interchangeably with ‘transient fecundity’ as they represent different life‐history parameters; (ii) provide a generalized re‐definition of the concept of fecundity selection as a mechanism that encompasses any traits that influence fecundity in any direction (from high to low) and in either sex; (iii) review the (macro)ecological basis of fecundity selection (e.g. ecological pressures that influence predictable spatial variation in fecundity); (iv) suggest that most ecological theories of fecundity selection should be tested in organisms other than birds; (v) argue that the longstanding fecundity selection hypothesis of female‐biased sexual size dimorphism (SSD) has gained inconsistent support, that strong fecundity selection does not necessarily drive female‐biased SSD, and that this form of SSD can be driven by other selective pressures; and (vi) discuss cases in which fecundity selection operates on males. This conceptual analysis of the theory of fecundity selection promises to help illuminate one of the central components of fitness and its contribution to adaptive evolution.     

The effect of water stress on the life cycles of Erodium crinitum Carolin and Erodium cicutarium (L.) L'Hérit. ex Alton (Geraniaceae)

Abstract Erodium cicutarium (L.) L'Hérit. ex Aiton (Geraniaceae) from temperate Mediterranean Eurasia is naturalized across large areas of arid and semi-arid Australia to which Erodium crinitum Carolin is native. plant size, leaf and bud numbers and fruit/plant biomass ratio were significantly lower under drought, flower and fruit number, fruit size and total mass were unaffected. In contrast, E. crinitum was largely unaffected by the drought conditions, showing only an increase in the fruit/plant biomass ratio.     

Demography and Life Histories of Sympatric Patas Monkeys, <Emphasis Type="Italic">Erythrocebus patas</Emphasis>, and Vervets, <Emphasis Type="Italic">Cercopithecus aethiops</Emphasis>, in Laikipia,Kenya

Mortality patterns are thought to be strong selective forces on life history traits, with high adult mortality and low immature mortality favoring early and rapid reproduction. Patas monkeys (Erythrocebus patas) have the highest potential rates of population increase for their body size of any haplorhine primate because they reproduce both earlier and more often. We report here 10 yr of comparative demographic data on a population of patas monkeys and a sympatric population of vervet monkeys (Cercopithecus aethiops), a closely related species differing in aspects of social system, ecology, and life history. The data reveal that 1) adult female patas monkeys have significantly higher mortality than adult female vervets; 2) infant mortality in patas monkeys is relatively low compared to the norm for mammals because it is not significantly different from that of adult female patas monkeys; and 3) infant mortality is significantly higher than adult female mortality in vervets. For both species, much of the mortality could be attributed to predation. An epidemic illness was also a major contributor to the mortality of adult female patas monkeys whereas chronic exposure to pathogens in a cold and damp microenvironment may have contributed to the mortality of infant vervets. Both populations experienced large fluctuations during the study period. Our results support the prediction from demographic models of life history evolution that high adult mortality relative to immature mortality selects for early maturation.     

One strategy does not fit all: determinants of urban adaptation in mammals

Urbanisation exposes wildlife to new challenging conditions and environmental pressures. Some mammalian species have adapted to these novel environments, but it remains unclear which characteristics allow them to persist. To address this question, we identified 190 mammals regularly recorded in urban settlements worldwide, and used phylogenetic path analysis to test hypotheses regarding which behavioural, ecological and life history traits favour adaptation to urban environments for different mammalian groups. Our results show that all urban mammals produce larger litters; whereas other traits such as body size, behavioural plasticity and diet diversity were important for some but not all taxonomic groups. This variation highlights the idiosyncrasies of the urban adaptation process and likely reflects the diversity of ecological niches and roles mammals can play. Our study contributes towards a better understanding of mammal association to humans, which will ultimately allow the design of wildlife‐friendly urban environments and contribute to mitigate human‐wildlife conflicts.