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Field biology of the platypus (Ornithorhynchus anatinus): historical and current perspectives.

The field biology of the platypus, Ornithorhynchus anatinus, was first studied by a number of expatriate biologists who visited the Australian colonies to collect specimens in the 1800s. Their work was followed in the early to mid-1900s by a group of resident natural historians and later by an increasing number of academic biologists. All of these workers contributed significantly to the current understanding of the field biology of this unique Australian species. The platypus occupies much the same general distribution as it did prior to European occupation of Australia, except for its loss from the state of South Australia. However, local changes and fragmentation of distribution due to human modification of its habitat are documented. The species currently inhabits eastern Australia from around Cooktown in the north to Tasmania in the south. Although not found in the west-flowing rivers of northern Queensland, it inhabits the upper reaches of rivers flowing to the west and north of the dividing ranges in the south of the state and in New South Wales and Victoria. Its current and historical abundance, however, is less well known and it has probably declined in numbers, although still being considered as common over most of its current range. The species was extensively hunted for its fur until around this turn of this century. The platypus is mostly nocturnal in its foraging activities, being predominantly an opportunistic carnivore of benthic invertebrates. The species is homeothermic, maintaining its low body temperature (32 degrees C), even while foraging for hours in water below 5 degrees C. Its major habitat requirements include both riverine and riparian features which maintain a supply of benthic prey species and consolidated banks into which resting and nesting burrows can be excavated. The species exhibits a single breeding season, with mating occurring in late winter or spring and young first emerging into the water after 3-4 months of nurture by the lactating females in the nesting burrows. Natural history observations, mark and recapture studies and preliminary investigations of population genetics indicate the possibility of resident and transient members of populations and suggest a polygynous mating system. Recent field studies have largely confirmed and extended the work of the early biologists and natural historians.

Animals↗

Intrafamilial conflict and parental investment: a synthesis.

We outline and develop current theory on how inherent genetic conflicts of interest between the various family members can affect the flow of parental investment from parents to offspring, and discuss the problems for empirical testing that this generates. The parental investment pattern realized in nature reflects the simultaneous resolution of all the conflicts between the family players. This depends on the genetic mechanism, the mating system and reproductive constraints, on whether extra demand by progeny affects current or future sibs, and particularly on the behavioural mechanisms underlying demand (begging or solicitation) and supply (provision of parental investment by parents). The direction of deviation from the optimal parental investment for the parent(s) depends on the slope of what we term the 'effect of supply on demand', the mechanism that determines how changes in food supply affect begging levels. If increasing food increases begging (positive slope), less parental investment is supplied than the parental optimum and if increasing food decreases begging (negative slope), more parental investment is supplied. The magnitude of deviation depends on both the 'effect of supply on demand' and on the 'effect of demand on supply' (the mechanism determining how changes in begging affect food supply, which always has a positive slope). We conclude that it will often be impossible to deduce the extent of underlying conflict by establishing the amount of parental investment given relative to the ideal optimum for the parent. Some possible directions for future research are discussed.

Animals↗

Pollen performance before and during the autotrophic-heterotrophic transition of pollen tube growth.

For species with bicellular pollen, the attrition of pollen tubes is often greatest where the style narrows at the transition between stigmatic tissue and the transmitting tissue of the style. In this region, the tubes switch from predominantly autotrophic to predominantly heterotrophic growth, the generative cell divides, the first callose plugs are produced, and, in species with RNase-type self-incompatibility (SI), incompatible tubes are arrested. We review the literature and present new findings concerning the genetic, environmental and stylar influences on the performance of pollen before and during the autotrophic-heterotrophic transition of pollen tube growth. We found that the ability of the paternal sporophyte to provision its pollen during development significantly influences pollen performance during the autotrophic growth phase. Consequently, under conditions of pollen competition, pollen selection during the autotrophic phase is acting on the phenotype of the paternal sporophyte. In a field experiment, using Cucurbita pepo, we found broad-sense heritable variation for herbivore-pathogen resistance, and that the most resistant families produced larger and better performing pollen when the paternal sporophytes were not protected by insecticides, indicating that selection during the autotrophic phase can act on traits that are not expressed by the microgametophyte. In a study of a weedy SI species, Solanum carolinense, we found that the ability of the styles to arrest self-pollen tubes at the autotrophic-heterotrophic transition changes with floral age and the presence of developing fruits. These findings have important implications for selection at the level of the microgametophyte and the evolution of mating systems of plants.

Cucurbita↗

Recent approaches into the genetic basis of inbreeding depression in plants.

Predictions for the evolution of mating systems and genetic load vary, depending on the genetic basis of inbreeding depression (dominance versus overdominance, epistasis and the relative frequencies of genes of large and small effect). A distinction between the dominance and overdominance hypotheses is that deleterious recessive mutations should be purged in inbreeding populations. Comparative studies of populations differing in their level of inbreeding and experimental approaches that allow selection among inbred lines support this prediction. More direct biometric approaches provide strong support for the importance of partly recessive deleterious alleles. Investigators using molecular markers to study quantitative trait loci (QTL) often find support for overdominance, though pseudo-overdominance (deleterious alleles linked in repulsion) may bias this perception. QTL and biometric studies of inbred lines often find evidence for epistasis, which may also contribute to the perception of overdominance, though this may be because of the divergent lines initially crossed in QTL studies. Studies of marker segregation distortion commonly uncover genes of major effect on viability, but these have only minor contributions to inbreeding depression. Although considerable progress has been made in understanding the genetic basis of inbreeding depression, we feel that all three aspects merit more study in natural plant populations.

Epistasis, Genetic↗

Genomic resources for comparative analyses of obligate avian brood parasitism.

Examples of convergent evolution, wherein distantly related organisms evolve similar traits, including behaviors, underscore the adaptive power of natural selection. In birds, obligate brood parasitism, and the associated loss of parental care behaviors, has evolved independently in seven different lineages, though little is known about the genetic basis of the complex suite of traits associated with this rare life history strategy. We generated genome assemblies for ten brood parasitic species plus eight species representatives of their parental/nesting outgroups. This includes nine long-read chromosome-level assemblies, with scaffold N50 sizes ranging from 38.1 to 72.6 MB, and gene representation completeness measures >97%. Leveraging this new catalog of avian genomes, we constructed clade-level alignments that reveal variation in chromosomal synteny, provide first-time or improved annotations of protein-coding and non-coding genes, and define cross-species ortholog reference sets. We also refine estimates for the timing of the seven independent origins of brood parasitism, ranging from recent events such as 1.6 to 4.5 million years ago in Molothrus cowbirds to much earlier origins over 30 million years ago in two of the three cuckoo lineages. These genomic resources lay the foundation for investigating the genetic and genomic underpinnings of brood parasitism, including the loss of parental care, shifts in mating systems, perhaps resulting in heightened sperm competition, elevated annual fecundity, improved spatial cognition related to nest-finding, and the diverse adaptations shaped by intense coevolution with host species.

assemblies↗

Frequency and spatial patterning of clonal reproduction in Louisiana iris hybrid populations.

The plant genera in which natural hybridization is most prevalent tend to be outcrossing perennials with some mechanism for clonal (i.e., asexual) reproduction. Although clonal reproduction in fertile, sexually reproducing hybrid populations could have important evolutionary consequences, little attention has been paid to quantifying this parameter in such populations. In the present study, we examined the frequency and spatial patterning of clonal reproduction in two Louisiana iris hybrid populations. Allozyme analysis of both populations revealed relatively high levels of genotypic diversity. However, a considerable amount of clonality was apparent. Nearly half of all genets (47%) in one population and more than half (61%) in the other had multiple ramets. Furthermore, both populations exhibited relatively high levels of genetic structuring, a pattern that resulted from the aggregation of clonal ramets. The occurrence of clonal reproduction in hybrid populations could not only facilitate introgression through an increase in the number of flowering ramets per genet and/or the survivorship of early generation hybrids, but might also influence the mating system of such populations. Any potential increase in the selfing rate due to cross-pollination among ramets of the same genet may, in turn, increase the likelihood of homoploid hybrid speciation.

Biological Evolution↗

Microevolutionary patterns and processes of the Native Hawaiian colonizing fern Odontosoria chinensis (Lindsaeaceae).

The vascular-plant flora of the Hawaiian Islands is characterized by one of the highest rates of species endemism in the world. Among flowering plants, approximately 89% of species are endemic, and among pteridophytes, about 76% are endemic. At the single-island level, however, rates of species endemism vary dramatically between these two groups with 80% of angiosperms and only 6% of pteridophytes being single-island endemics. Thus, in many groups of Hawaiian angiosperms, it is possible to link studies of phylogeny, evolution, and biogeographic history at the interspecific and interisland levels. In contrast, the low level of single-island species endemism among Hawaiian pteridophytes makes similar interspecific and interisland studies nearly impossible. Higher levels of interisland gene flow may account for the different levels of single-island endemism in Hawaiian pteridophytes relative to angiosperms. The primary question we addressed in the present study was: Can we infer microevolutionary patterns and processes among populations within widespread species of Hawaiian pteridophytes wherein gene flow is probably common? To address this broad question, we conducted a population genetic study of the native Hawaiian colonizing species Odontosoria chinensis. Data from allozyme analyses allowed us to infer: (1) significant genetic differentiation among populations from different islands; (2) historical patterns of dispersal between particular pairs of islands; (3) archipelago-level patterns of dispersal and colonization; (4) founder effects among populations on the youngest island of Hawaii; and, (5) that this species primarily reproduces via outcrossing, but may possess a mixed-mating system.

Biological Evolution↗

Phylogenetic analysis of sexual dimorphism and eye-span allometry in stalk-eyed flies (Diopsidae).

Eye stalks and their scaling relationship with body size are important features in the mating system of many diopsid species, and sexual selection is a critical force influencing the evolution of this exaggerated morphology. Interspecific variation in eye span suggests there has been significant evolutionary change in this trait, but a robust phylogenetic hypothesis is required to determine its rate and direction of change. In this study, the pattern of morphological evolution of eye span is assessed in a phylogenetic framework with respect to its function in the sexual system of these flies. Specifically, we examine within the family Diopsidae the pattern of increase and decrease in sexual dimorphism, the morphological coevolution of eye span between males and females, and the evolutionary flexibility of eye-span allometry. Based on several different methods for reconstructing morphological change, results suggest a general pattern of evolutionary flexibility, particularly for eye-span allometry. Sexual dimorphism in eye span has evolved independently at least four times in the family and this trait also has undergone several reductions within the genus Diasemopsis. Despite most species being dimorphic, there is a strong phylogenetic correlation between males and females for mean eye span. The coevolution between the sexes for eye-span allometry, however, is significantly weaker. Overall, eye-span allometry exhibits significantly more change on the phylogeny than the other morphological traits. The evolutionary pattern in eye-span allometry is caused primarily by changes in eye-span variance. Therefore, this pattern is consistent with recent models that predict a strong relationship between sexual selection and the variance of ornamental traits and highlights the significance of eye-span allometry in intersexual and intrasexual signaling.

Animals↗

Hybrid bridges to gene flow: a case study in milkweeds (Asclepias).

Natural hybridization occurs throughout areas of sympatry for the North American milkweeds Asclepias exaltata and A. syriaca (Asclepiadaceae), even though the formation of F1 hybrid seed is a rare event. For introgressive hybridization to proceed, F1 and advanced hybrids must be released from reproductive barriers and successfully mate with one or both parental species. I investigated the mating system of natural hybrids between A. exaltata and A. syriaca in three populations in Shenandoah National Park, Virginia. Allozyme data and a maximum-likelihood procedure were used to estimate the frequency of six genotypic classes (parentals, F1, F2, and backcrosses) of the hybridizing populations, the pollinia received by hybrid plants, and the paternal parents of seeds produced by hybrids. F1 hybrids, backcross A. syriaca, and parental A. syriaca individuals were common in three hybrid populations. Even though self-pollinations and interhybrid pollinations were common, F2 seed production and the occurrence of F2 individuals were rare in hybrid populations. Hybrid plants received more pollen from A. syriaca than A. exaltata, which resulted in the production of more backcross-A. syriaca seed than backcross-A. exaltata seed. Asclepias exaltata was rare in the hybrid populations, but A. exaltata pollinia were received by hybrids and this species sired between 15% and 36% of the seeds produced on hybrids. The potential for introgression with A. exaltata populations is lower because this species is unsuccessful as the maternal parent in interspecific and backcross hand-pollinations. The asymetry of hybridization with A. syriaca as the maternal parent is further supported by the incorporation of maternally inherited chloroplast DNA markers in hybrids. Hybrid milkweeds frequently backcross with both parental species and may be released from the reproductive barriers that limit the formation of F1 hybrids in natural populations. The direction of interspecific gene flow and introgression in milkweeds is influenced by the reproductive biology of hybrids, the constituency of the surrounding population, and failure of some crosses to produce seeds. Finally, introgressive hybridization remains an important evolutionary force even when the initial formation of F1 hybrids in natural populations is rare.

Alleles↗

A metapopulation perspective on genetic diversity and differentiation in partially self-fertilizing plants.

Partial self-fertilization is common in higher plants. Mating system variation is known to have important consequences for how genetic variation is distributed within and among populations. Selfing is known to reduce effective population size, and inbreeding species are therefore expected to have lower levels of genetic variation than comparable outcrossing taxa. However, several recent empirical studies have shown that reductions in genetic diversity within populations of inbreeding species are far greater than the expected reductions based on the reduced effective population size. Two different processes have been argued to cause these patterns, selective sweeps (or hitchhiking) and background selection. Both are expected to be most effective in reducing genetic variation in regions of low recombination rates. Selfing is known to reduce the effective recombination rate, and inbreeding taxa are thus thought to be particularly vulnerable to the effects of hitchhiking or background selection. Here I propose a third explanation for the lower-than-expected levels of genetic diversity within populations of selfing species; recurrent extinctions and recolonizations of local populations, also known as metapopulation dynamics. I show that selfing in a metapopulation setting can result in large reductions in genetic diversity within populations, far greater than expected based the lower effective population size inbreeding species is expected to have. The reason for this depends on an interaction between selfing and pollen migration.

Genetic Variation↗

Species divergence in sexually selected traits: increase in song elaboration is related to decrease in plumage ornamentation in finches.

Elaborate plumage and complex songs of male birds are two of the best-known examples of sexually selected traits, yet the interaction between these traits is poorly understood. Theory suggests that among a suite of potential displays, animals will emphasize traits that are most conspicuous, least costly, or best signal condition and reduce display of other traits. Here we examined the relationship between song and plumage elaborations in cardueline finches, songbirds that are highly variable in plumage displays and songs, but that share a similar mating system. We statistically controlled for body mass, habitat characteristics, and phylogenetic relationships and found that across species song complexity was strongly negatively related to elaboration of plumage ornamentation. When plumage coloration was partitioned into carotenoid-based and melanin-based components, song complexity was negatively related to elaboration of male carotenoid-based coloration but unrelated to elaboration of melanin-based coloration. These observations support the idea that, for condition-dependent traits, animal species trade off trait expression in response to changes in the costs or the information content of these traits. We discuss several alternative explanations for the observed pattern.

Animals↗

The origins of sexual dimorphism in body size in ungulates.

Jarman (1974) proposed a series of relationships between habitat use, food dispersion, and social behavior and hypothesized a series of evolutionary steps leading to sexual dimorphism in body size through sexual selection in African antelope species. The hypothesis states that sexual size dimorphism evolved in a three-step process. Initially, ancestral monomorphic and monogamous ungulate species occupying closed habitats radiated into open grassland habitats. Polygynous mating systems then rapidly evolved in response to the aggregation of males and females, perhaps in relation to the clumped distribution of food resources in open habitats. Subsequently, size dimorphism evolved in those species occupying open habitats, but not in species that remained in closed habitats or retained monogamy. This hypothesis has played an important role in explaining the origins of sexual dimorphism in mammals. However, the temporal sequence of the events that Jarman proposed has never been demonstrated. Here we use a phylogeny of extant ungulate species, along with maximum-likelihood statistical techniques, to provide a test of Jarman's hypothesis.

Animals↗

Inbreeding and sex-biased gene flow in the ant Formica exsecta.

The objective of this study was to assess breeding and dispersal patterns of both males and females in a monogyne (a single queen per colony) population of ants. Monogyny is commonly associated with extensive nuptial flights, presumably leading to considerable gene flow over large areas. Opposite to these expectations we found evidence of both inbreeding and sex-biased gene flow in a monogyne population of Formica exsecta. We found a significant degree of population subdivision at a local scale (within islands) for queens (females heading established colonies) and workers, but not for colony fathers (the males mated to the colony queens). However, we found little evidence of population subdivision at a larger scale (among islands). More conclusive support for sex-biased gene flow comes from the analysis of isolation by distance on the largest island, and from assignment tests revealing differences in female and male philopatry. The genetic similarity between pairs of queens decreased significantly when geographical distance increased, demonstrating limited dispersal and isolation by distance in queens. By contrast, we found no such pattern for colony fathers. Furthermore, a significantly greater fraction of colony queens were assigned as having originated from the population of residence, as compared to colony fathers. Inbreeding coefficients were significantly positive for workers, but not for mother queens. The queen-male relatedness coefficient of 0.23 (regression relatedness) indicates that mating occurs between fairly close relatives. These results suggest that some monogyne species of ants have complex dispersal and mating systems that can result in genetic isolation by distance over small geographical scales. More generally, this study also highlights the importance of identifying the relevant scale in analyses of population structure and dispersal.

Animals↗

Viability selection at three early life stages of the tropical tree, Platypodium elegans (Fabaceae, Papilionoideae).

Given the enormous number and high mortality of fertilized ovules in plants, it is possible that selection during the earliest stages of the life cycle plays an important role in shaping the genetic composition of plant populations. Previous research involving selection component analyses found strong evidence for viability selection in annual plant species. Yet despite this evidence, few attempts have been made to identify the magnitude and timing of viability selection as well as the mechanisms responsible for mortality among genotypes. Platypodium elegans, a Neotropical tree with high rates of early fruit mortality, represents an opportunity to study viability selection at a level of discernment not previously possible. Microsatellite markers were used to analyze the genetic composition of aborted embryos, as well as mature seeds and seedlings of the same cohort. While selection resulted in an overall decrease in self-fertilized progeny across each life stage, the greatest change in the genetic composition of progeny occurred between mature seeds and established seedlings. This suggested that inbreeding depression, and not late-acting self-incompatibility, was responsible for early selection. An investigation of the mature seed stage revealed that self-fertilized seeds weigh significantly less than outcrossed seeds. The result of this early selection conceals the mixed-mating system and high levels of inbreeding depression in Platypodium elegans, resulting in an apparently outcrossed adult population that does not differ significantly from Hardy-Weinberg expectations.

Alleles↗

Mechanisms of sperm competition: testing the fair raffle.

Sperm competition is a major force of sexual selection, but its implications for mating system and life-history evolution are just beginning to be understood. Of particular importance is understanding the mechanisms of sperm competition. Models have been developed to determine if sperm competition operates in a fair raffle process, whereby each sperm from competing males has an equal chance of fertilizing a female's ova, or if it operates in a loaded raffle process, whereby one male's sperm has a fertilization advantage. These models require data on relative sperm and offspring (paternity) numbers of competing males. Here we develop a model based on maximum-likelihood methods for differentiating between the fair and loaded raffle processes. The model calculates the relative competitiveness of two males' sperm (loadings) as well as the economy of scale (nonlinear returns to sperm number). Previous models implicitly assumed that there is no economy of scale, which may not be the case when there is cooperation or interference among sperm from a given male. We demonstrate that our model has superior power-in some instances more than double-than previous models. We apply our model to an example of sperm competition in the guppy (Poecilia reticulata) and show that the system may be characterized by a loaded raffle attributable to effects of second male precedence.

Animals↗

Heterozygote excess in small populations and the heterozygote-excess effective population size.

It has been proposed that effective size could be estimated in small dioecious population by considering the heterozygote excess observed at neutral markers. When the number of breeders is small, allelic frequencies in males and females will slightly differ due to binomial sampling error. However, this excess of heterozygotes is not generated by dioecy but by the absence of individuals produced through selfing. Consequently, the approach can also be applied to self-incompatible monoecious species. Some inaccuracies in earlier equations expressing effective size as function of the heterozygote excess are also corrected in this paper. The approach is then extended to subdivided populations, where time of sampling becomes crucial. When adults are sampled, the effective size of the entire population can be estimated, whereas when juveniles are sampled, the average effective number of breeders per subpopulations can be estimated. The main limitation of the heterozygote excess method is that it will only perform satisfactorily for populations with a small number of reproducing individuals. While this situation is unlikely to happen frequently at the scale of the entire population, structured populations with small subpopulations are likely to be common. The estimation of the average number of breeders per subpopulations is thus expected to be applicable to many natural populations. The approach is straightforward to compute and independent of equilibrium assumptions. Applications to simulated data suggest the estimation of the number of breeders to be robust to mutation and migration rates, and to specificities of the mating system.

Biological Evolution↗

A comparative study of the bond strengths of amalgam and alloy-glass ionomer cores.

The purpose of this study was to compare the bond strengths of dental amalgam cores and dental alloy-glass ionomer cores that were luted to cast gold crowns with glass ionomer cement. Seventy-two human extracted molars were sectioned horizontally and four regular thread mate system pins were inserted into a flat pulpal floor. The teeth were restored with amalgam or alloy-glass ionomer admixture and prepared for crown preparations. Castings were produced with type III gold and cemented to the cores with Fuji type I glass ionomer cement. Thirty-six of the specimens were subjected to thermal fatigue by cycling between 4 and 50 degrees C for periods of 1, 5 and 10 weeks. The remainder of the specimens were retained in deionized water at 37 degrees C. Bond strengths of the cores, in tension, were measured with a universal testing machine. The alloy-glass ionomer cores exhibited higher bond strengths than the amalgam cores for virtually all time periods, particularly the thermocycled samples. The alloy-glass ionomer cores exclusively displayed core fractures and pin/tooth insufficiencies while the amalgam cores failed because of a deterioration of the glass ionomer luting cement.

Crowns↗

Deterministic paternity exclusion using RAPD markers.

The Random Amplified Polymorphic DNA (RAPD) technique can potentially provide hundreds of polymorphic markers for use by ecologists studying mating systems in natural populations. We consider here the implications of the dominance displayed by RAPD markers for deterministic paternity assignment. Our goal was to provide a means for assessing the costs associated with such a study for ecologists who might be considering the use of RAPD markers for paternity analysis. The theoretical expected proportion of offspring for which all males except the true father can be exlucded (P(ET)) is calculated for both dominant and codominant marker systems. The ability to assign paternity unambiguously generally increases with the number of loci and the frequency of the recessive allele (but only up to a point), and decreases with increasing sample size (number of individuals surveyed). The gain in P(ET) with decreasing sample size is unexpectedly slight. Not surprisingly, the performance of dominant markers at paternity exclusion is, in general, greatly exceeded by codominant markers, with the exception of the case in which the frequency of the recessive allele is high at all loci. In this case, codominant markers perform only slightly better than do dominant markers. Thus, a researcher should expect to score more than 50 RAPD loci for each offspring for most applications of paternity exclusion analysis.(ABSTRACT TRUNCATED AT 250 WORDS)

Alleles↗