Coevolution of male and female genital morphology in waterfowl

Most birds have simple genitalia; males lack external genitalia and females have simple vaginas. However, male waterfowl have a phallus whose length (1.5->40 cm) and morphological elaborations vary among species and are positively correlated with the frequency of forced extra-pair copulations among waterfowl species. Here we report morphological complexity in female genital morphology in waterfowl and describe variation vaginal morphology that is unprecedented in birds. This variation comprises two anatomical novelties: (i) dead end sacs, and (ii) clockwise coils. These vaginal structures appear to function to exclude the intromission of the counter-clockwise spiralling male phallus without female cooperation. A phylogenetically controlled comparative analysis of 16 waterfowl species shows that the degree of vaginal elaboration is positively correlated with phallus length, demonstrating that female morphological complexity has co-evolved with male phallus length. Intersexual selection is most likely responsible for the observed coevolution, although identifying the specific mechanism is difficult. Our results suggest that females have evolved a cryptic anatomical mechanism of choice in response to forced extra-pair copulations.     

Coevolution of parasite virulence and host life history

Most models about the evolutionary interactions between a parasite's virulence and its host's life history neglect two potentially important aspects: epidemiological and coevolutionary feedback. We emphasize their importance by presenting models that describe the coevolution of a semelparous host's age at reproduction and a parasite's virulence in different environmental conditions. In particular, we first show that an epidemiological feedback will lead to a nonmonotonic response of the host's age at reproduction as virulence increases. We then show that the coevolutionary pressure on virulence can lead to complex associations between the host's life history and the parasite's virulence, which would not be expected with more traditional models of host or parasite evolution. Thus, for example, a high mortality rate of the host favours avirulent parasites and late reproduction of the host when the environmental conditions allow the host to grow rapidly, but early reproduction and high virulence when growth is slow.     

Coevolution of sperm and female reproductive tract morphology in stalk-eyed flies

Sperm and female reproductive tract morphology are among the most rapidly evolving characters known in insects. To investigate whether interspecific variation in these traits results from divergent coevolution we examined testis size, sperm length and female reproductive tract morphology for evidence of correlated evolution using 13 species of diopsid stalk-eyed flies. We found that sperm dimorphism (the simultaneous production of two size classes of sperm by individual males) is ancestral and occurs in four genera while sperm monomorphism evolved once and persists in one genus. The length of ''long-sperm'' types, though unrelated to male body or testis size, exhibits correlated evolution with two regions of the female reproductive tract, the spermathecae and ventral receptacle, where sperm are typically stored and used for fertilization, respectively. Two lines of evidence indicate that ''short sperm'', which are probably incapable of fertilization, coevolve with spermathecae. First, loss of sperm dimorphism coincides phylogenetically with reduction or loss of spermathecae. Second, evolutionary change in short-sperm length correlates with change in spermathecal size but not spermathecal duct length or ventral receptacle length. Morphological coevolution between sperm and female reproductive tracts is consistent with a history of female-mediated selection on sperm length.     

Pattern and process in the biogeography of subterranean amphipods

The comparative data from studies of two ‘superfamily’ groups with large numbers of subterranean taxa, the exclusively freshwater Crangonyctoidea and the predominately marine Hadzioidea, support the hypothesis that distributional patterns and evolutionary processes of stygobiont amphipods are closely linked and that the former can be a useful indicator of the latter. Three major biogeographic patterns are indicated by the distribution of subterranean species in these groups, each apparently reflecting a particular mode of origin: (1) freshwater stygobionts (limnostygobionts) derived from epigean freshwater ancestors through colonizations probably influenced by adaptive shifts, or assisted by stream capture and spring failure; (2) freshwater stygobionts derived from marine/brackish water ancestors by stranding during regression of marine embayments; and (3) marine/brackish water stygobionts (thalassostygobionts) derived from epigean marine/brackish water ancestors through adaptive shifts possibly in concert with fluctuating sea levels.     

Coevolution of male and female reproductive traits in a simultaneously hermaphroditic land snail

Inter- and intraspecific studies in gonochoristic animals reveal a covariation between sperm characteristics and the size of the female reproductive tract, indicating a rapid evolutionary divergence, which is consistent with the theory of post-copulatory sexual selection. Simultaneous hermaphrodites differ from species with separate sexes (gonochorists) in that they possess both functional male and female reproductive organs at the same time. We investigated whether in hermaphroditic animals intraspecific variation in reproductive traits results from divergent coevolution, by quantifying the variation in male and female traits among six natural populations of the snail Arianta arbustorum and examining the covariation in interacting traits. There was a significant among-population variation in spermatophore volume, number of sperm transferred and sperm length, as well as in volume of the sperm storage organ (spermatheca) and number of tubules, but not in spermatheca length. We found a positive association between sperm number transferred and spermatheca volume. This result suggests that the same post-copulatory mechanisms as in gonochorists drive the correlated evolution of reproductive characters in hermaphrodites.     

Foraging strategies are related to skull morphology and life history traits of Melanerpes woodpeckers

Links between morphology and foraging strategies have been well established for many vertebrate groups. Foraging strategies of Melanerpes woodpeckers are especially variable, with at least six species being proficient flycatchers; the remainder of the better known species do not flycatch. Our objective was to examine variation in foraging tactics as it relates to skull morphology and other life history traits among these species to better understand the biology of these diverse woodpeckers. We measured eight skull characters from 241 individuals representing 19 species, but focused on eight species for which we had the most data. We used the log-geometric mean and a principal components analysis (PCA) to calculate size-scaled shape variables. Cluster analysis based on PCA scores clearly separated birds by foraging behavior. Species with similar foraging behaviors (i.e., flycatchers vs non-flycatchers) also share a number of other life history characteristics including similar plumage, diets, and migratory behavior. Diversity within Melanerpes may imply a high degree of plasticity or that species have been incorrectly placed in a polyphyletic group. Woodpeckers currently in the genus Melanerpes share few uniting characters and historically have been placed in as many as eight different genera. Additional life history, morphological, and genetic studies of the group, especially of Caribbean and Neotropical species, is warranted.     

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.     

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.     

A review of the reproductive bionomics of aquatic gammaridean amphipods: variation of life history traits with latitude,depth, salinity and superfamily

Life history traits (mean and maximum body length of females, number of embryos per brood = brood size, embryo diameter, number of broods per female, lifespan of females) for 302 populations of aquatic gammaridean amphipods, representing 214 species in 16 superfamilies, were reviewed. The variation of these traits, of lifetime potential fecundity (i.e. the number of embryos produced per female lifespan) and of reproductive potential (i.e. the number of embryos produced per female per year), with temperature (latitude), depth, salinity and superfamily, was investigated by various univariate and multivariate methods. Gammaridean amphipods comprise semelparous and iteroparous populations and species, with semiannual, annual, biannual or perennial life cycles. However, most gammarideans studied so far are iteroparous annuals. Body length explains most of the variation in brood size and embryo diameter. The reproductive potential may be increased by increasing body size for a constant breeding frequency, by increasing brood size at the expense of smaller embryos, by increasing breeding frequency for a constant lifespan at the expense of smaller individual broods and/or embryos, and by increasing longevity for a constant breeding frequency and brood size. Combinations of these different options constitute the life history patterns of gammarideans, which vary across superfamilies, latitude and depth, and cannot simply be explained by variations in body length. High latitude species were generally characterized by biannual or perennial life histories, large body size, delayed maturity, and single or few broods with many, relatively large embryos; converse sets of traits characterized low latitude species. Deep-living species had relatively smaller broods and embryos than their shallow-living relatives, yet did not produce more broods. However, different superfamilies dominated in different habitats. The importance of natural selection relative to phylogenetic (historical) and physiological constraints in the forging of these patterns is discussed.     

Cost of reproduction and covariation of life history traits in birds

Evolution of life history traits can be studied at two different levels: (1) current selection processes, including trade-offs in life history traits in natural populations as revealed by observations or, preferably, exieriments; and (2) patterns of variation in life history traits with each other and with ecology among extant species. Selection is not evolution, but selection pressures must have caused evolutionary change and led to current patterns of life history traits. These problems are exemplified by recent research on clutch size in birds.     

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.     

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.     

Coevolution between parasite virulence and host life-history traits

Epidemiological models generally explore the evolution of parasite life-history traits, namely, virulence and transmission, against a background of constant host life-history traits. However, life-history models have predicted the evolution of host traits in response to parasitism. The coevolution of host and parasite life-history traits remains largely unexplored. We present an epidemiological model, based on resource allocation theory, that provides an analysis of the coevolution between host reproductive effort and parasite virulence. This model allows for hosts with either a fixed (i.e., genetic) or conditional (i.e., a phenotypically plastic) response to parasitism. It also considers superinfections. We show that parasitism always favors increased allocation to host reproduction, but because of epidemiological feedbacks, the evolutionarily stable host reproductive effort does not always increase with parasite virulence. Superinfection drives the evolution of parasite virulence and acts on the evolution of the host through parasite evolution, generally leading to higher host reproductive effort. Coevolution, as opposed to cases where only one of the antagonists evolves, may generate correlations between host and parasite life-history traits across environmental gradients affecting the fecundity or the survival of the host. Our results provide a theoretical framework against which experimental coevolution outcomes or field observations can be contrasted.     

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.     

Coevolution of male and female reproductive structures in Drosophila

The morphology of male genitalia whilst stable within species, exhibits huge interspecific variation. This variation is likely to be as a result of sexual selection due to the direct involvement of these reproductive structures in mating and sperm transfer. In contrast, internal soft tissue components of the genitalia are generally poorly investigated as they are not directly involved in physical and mechanical adequacy during sperm transfer. However, these soft tissue structures may also drive differential male–female interactions, particularly in internally fertilising organisms where females have the ability to store sperm and bias male reproductive success. In this paper we use the drosophila model to investigate the role of male and female reproductive elements in sexual selection. Our meta-analysis supplemented with additional new data clearly shows that within species, sperm length versus testis length, and sperm length versus seminal receptacle length, are highly correlated. Thus, independent of the phylogenetic relationship among species, gamete evolution is likely to result in sexual selection interactions that drive the evolution of internal reproductive components in both sexes. Our results and discussion of the literature highlight the importance of considering internal soft structures that may influence fertilisation, when investigating selective forces acting on the evolution of reproductive traits.     

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.