Variation In Reproduction Research Articles

Rampant asexual reproduction and limited dispersal in a mangrove population of the coral Porites divaricata.

Corals are critical to marine biodiversity. Reproduction and dispersal are key to their resilience, but rarely quantified in nature. Exploiting a unique system-a fully censused, longitudinally characterized, semi-isolated population inhabiting mangroves-we used 2bRAD sequencing to demonstrate that rampant asexual reproduction most likely via parthenogenesis and limited dispersal enable the persistence of a natural population of thin-finger coral (Porites divaricata). Unlike previous studies on coral dispersal, knowledge of colony age and location enabled us to identify plausible parent-offspring relationships within multiple clonal lineages and develop tightly constrained estimates of larval dispersal; the best-fitting model indicates dispersal is largely limited to a few metres from parent colonies. Our results explain why this species is adept at colonizing mangroves but suggest limited genetic diversity in mangrove populations and limited connectivity between mangroves and nearby reefs. As P. divaricata is gonochoristic, and parthenogenesis would be restricted to females (whereas fragmentation, which is presumably common in reef and seagrass habitats, is not), mangrove populations likely exhibit skewed sex ratios. These findings suggest that coral reproductive diversity can lead to distinctly different demographic outcomes in different habitats. Thus, coral conservation will require the protection of the entire coral habitat mosaic, and not just reefs.

Циклическое развитие проектно-технологической культуры: проблемы методологии

The article analyzes approaches to understanding the essence of technological culture and the patterns of its development from the methodological positions of the design culture and the theory of selforganization. The phenomenon of technological culture is studied in the unity of the conceptual, praxeological and organizational aspects. The process of formation and development of technological culture is cyclical. Cyclicity is appearing in the periodic reiteration of the stages of cultural reproduction and cultural variability. The study gives a description of the three-phase structure of the cultural cycle, taking into account the role of self-organization processes at each formative stage. At the first formative stage the specificity of the self-organization mechanism is reduced to the formation of rigid links (structures) that measure and fasten images, signs and objects that are useful for the cultural formation. At the second stage a fundamental livelihood of the developing object as a whole is created. Usually this is a fundamental principle that sets coherence, synchronization of processes in a sociocultural system. At the third stage the processes of self-organization provide a livelihood for the components of the object which leads to the achievement of morphological and axiological integrity and uniqueness of the object, the system enters into symbiotic relationships with objects of another type, completes development and prepares the transition to a new three-part cycle of shaping. The article deals with the first historical cycle of formation and development of technological culture: "Hellenism – Ancient Rome – Byzantium". Each phase of the cultural cycle is characterized by the originality of subject, target and organizational factors. In future the undertaken research will reveal the features of technological culture on its modern "digital" stage basing on the logic of cyclic patterns developed within the framework of the proposed methodology.

Assessment of management and reproductive variability among the buffalo farms in Chitwan based on the scores of production efficiency

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The effect of prey abundance and fisheries on the survival, reproduction, and social structure of killer whales (Orcinus orca) at subantarctic Marion Island.

Most marine apex predators are keystone species that fundamentally influence their ecosystems through cascading top-down processes. Reductions in worldwide predator abundances, attributed to environmental- and anthropogenic-induced changes to prey availability and negative interactions with fisheries, can have far-reaching ecosystem impacts. We tested whether the survival of killer whales (Orcinus orca) observed at Marion Island in the Southern Indian Ocean correlated with social structure and prey variables (direct measures of prey abundance, Patagonian toothfish fishery effort, and environmental proxies) using multistate models of capture-recapture data spanning 12 years (2006-2018). We also tested the effect of these same variables on killer whale social structure and reproduction measured over the same period. Indices of social structure had the strongest correlation with survival, with higher sociality associated with increased survival probability. Survival was also positively correlated with Patagonian toothfish fishing effort during the previous year, suggesting that fishery-linked resource availability is an important determinant of survival. No correlation between survival and environmental proxies of prey abundance was found. At-island prey availability influenced the social structure of Marion Island killer whales, but none of the variables explained variability in reproduction. Future increases in legal fishing activity may benefit this population of killer whales through the artificial provisioning of resources they provide.

Environmental and molecular regulation of asexual reproduction in the sea anemone Nematostella vectensis

Cnidarians exhibit incredible reproductive diversity, with most capable of sexual and asexual reproduction. Here, we investigate factors that influence asexual reproduction in the burrowing sea anemone Nematostella vectensis, which can propagate asexually by transverse fission of the body column. By altering culture conditions, we demonstrate that the presence of a burrowing substrate strongly promotes transverse fission. In addition, we show that animal size does not affect fission rates, and that the plane of fission is fixed along the oral–aboral axis of the polyp. Homeobox transcription factors and components of the TGF β , Notch, and FGF signalling pathways are differentially expressed in polyps undergoing physal pinching suggesting they are important regulators of transverse fission. Gene ontology analyses further suggest that during transverse fission the cell cycle is suppressed, and that cell adhesion and patterning mechanisms are downregulated to promote separation of the body column. Finally, we demonstrate that the rate of asexual reproduction is sensitive to population density. Collectively, these experiments provide a foundation for mechanistic studies of asexual reproduction in Nematostella, with implications for understanding the reproductive and regenerative biology of other cnidarian species.

Hurricanes affect diversification among individual life courses of a primate population.

Extreme climatic events may influence individual-level variability in phenotypes, survival and reproduction, and thereby drive the pace of evolution. Climate models predict increases in the frequency of intense hurricanes, but no study has measured their impact on individual life courses within animal populations. We used 45 years of demographic data of rhesus macaques to quantify the influence of major hurricanes on reproductive life courses using multiple metrics of dynamic heterogeneity accounting for life course variability and life-history trait variances. To reduce intraspecific competition, individuals may explore new reproductive stages during years of major hurricanes, resulting in higher temporal variation in reproductive trajectories. Alternatively, individuals may opt for a single optimal life-history strategy due to trade-offs between survival and reproduction. Our results show that heterogeneity in reproductive life courses increased by 4% during years of major hurricanes, despite a 2% reduction in the asymptotic growth rate due to an average decrease in mean fertility and survival by that is, shortened life courses and reduced reproductive output. In agreement with this, the population is expected to achieve stable population dynamics faster after being perturbed by a hurricane ( ; 95% CI: 1.488, 1.538), relative to ordinary years . Our work suggests that natural disasters force individuals into new demographic roles to potentially reduce competition during unfavourable environments where mean reproduction and survival are compromised. Variance in lifetime reproductive success and longevity are differently affected by hurricanes, and such variability is mostly driven by survival.

Five new species of the pelodryadid genus Litoria Tschudi from the southern versant of Papua New Guinea’s Central Cordillera, with observations on the diversification of reproductive strategies in Melanesian treefrogs

New Guinea has the most diverse insular frog fauna in the world, and rates of species discovery and description have increased rapidly in the last two decades. Pelodryadid treefrogs are the second most diverse family of anurans on the island but their taxonomy, relationships, and especially ecology remain poorly documented. Based on differences in morphology, advertisement calls (where available) and phylogenetic analyses of a 787 base pair alignment from the mitochondrial ND4 gene and flanking tRNA, we describe five new species of small treefrogs from hill and lower montane forests in the high rainfall belt that straddles the southern versant of Papua New Guinea's Central Cordillera. Three of these species are known only from forest growing on karst substrates, adding to the growing number of herpetofauna species currently known only from the extensive karst habitats of Papua New Guinea's South-fold Mountains. We also describe the arboreal breeding strategies of two of the new species, and report obligate treehole (phytotelm) breeding in New Guinean frogs for the first time. The new phytotelm-breeding species has juveniles with colour and patterning that closely resemble bird droppings, suggesting defensive mimicry or masquerade. A preliminary phylogeny suggests that arboreal-breeding frogs do not form a monophyletic group and that arboreal breeding has evolved multiple times within the New Guinean pelodryadid radiation. A further striking feature of the phylogeny is poor support for most basal nodes in the most diverse radiation of Melanesian Pelodryadidae, suggesting rapid ecological diversification and speciation, potentially following colonisation from Australia and/or mountain uplift. These new taxa and observations highlight previously unrecognised ecological and reproductive diversity in the Melanesian Pelodryadidae.

Reproductive variance can drive behavioral dynamics

The concept of fitness is central to evolution, but it quantifies only the expected number of offspring an individual will produce. The actual number of offspring is also subject to demographic stochasticity-that is, randomness associated with birth and death processes. In nature, individuals who are more fecund tend to have greater variance in their offspring number. Here, we develop a model for the evolution of two types competing in a population of nonconstant size. The fitness of each type is determined by pairwise interactions in a prisoner's dilemma game, and the variance in offspring number depends upon its mean. Although defectors are preferred by natural selection in classical population models, since they always have greater fitness than cooperators, we show that sufficiently large offspring variance can reverse the direction of evolution and favor cooperation. Large offspring variance produces qualitatively new dynamics for other types of social interactions, as well, which cannot arise in populations with a fixed size or with a Poisson offspring distribution.

An oligogenic architecture underlying ecological and reproductive divergence in sympatric populations

The evolutionary trajectories and genetic architectures underlying ecological divergence with gene flow are poorly understood. Sympatric timing types of the intertidal insect Clunio marinus (Diptera) from Roscoff (France) differ in lunar reproductive timing. One type reproduces at full moon, the other at new moon, controlled by a circalunar clock of yet unknown molecular nature. Lunar reproductive timing is a magic trait for a sympatric speciation process, as it is both ecologically relevant and entails assortative mating. Here, we show that the difference in reproductive timing is controlled by at least four quantitative trait loci (QTL) on three different chromosomes. They are partly associated with complex inversions, but differentiation of the inversion haplotypes cannot explain the different phenotypes. The most differentiated locus in the entire genome, with QTL support, is the period locus, implying that this gene could not only be involved in circadian timing but also in lunar timing. Our data indicate that magic traits can be based on an oligogenic architecture and can be maintained by selection on several unlinked loci.

Population structure and diversity of the needle pathogen Dothistroma pini suggests human-mediated movement in Europe.

Dothistroma needle blight (DNB) is an important disease of Pinus species that can be caused by one of two distinct but closely related pathogens; Dothistroma septosporum and Dothistroma pini. Dothistroma septosporum has a wide geographic distribution and is relatively well-known. In contrast, D. pini is known only from the United States and Europe, and there is a distinct lack of knowledge regarding its population structure and genetic diversity. The recent development of 16 microsatellite markers for D. pini provided an opportunity to investigate the diversity, structure, and mode of reproduction for populations collected over a period of 12years, on eight different hosts in Europe. In total, 345 isolates from Belgium, the Czech Republic, France, Hungary, Romania, Western Russia, Serbia, Slovakia, Slovenia, Spain, Switzerland, and Ukraine were screened using microsatellite and species-specific mating type markers. A total of 109 unique multilocus haplotypes were identified and structure analyses suggested that the populations are influenced by location rather than host species. Populations from France and Spain displayed the highest levels of genetic diversity followed by the population in Ukraine. Both mating types were detected in most countries, with the exception of Hungary, Russia and Slovenia. Evidence for sexual recombination was supported only in the population from Spain. The observed population structure and several shared haplotypes between non-bordering countries provides good evidence that the movement of D. pini in Europe has been strongly influenced by human activity in Europe.

Reproductive functional organization of woody plant assemblages along regeneration in a Caatinga dry forest

Tropical forest regeneration across old fields has been mainly described as a predictable sequence of functional plant assemblages in response to environmental filtering. However, the way plant reproductive diversity is organized along forest regeneration and how the reproductive profile of woody flora may impact regeneration have been poorly investigated. This paper examines the organization of plant assemblages throughout the regeneration of a Caatinga dry forest in the context of slash-and-burn agriculture with a focus on reproductive plant traits and strategies. We hypothesized a recovery and directional changes in specialist reproductive traits/strategies along forest regeneration but with aridity and chronic anthropogenic disturbances posing contrary forces to this recovery process. This study was carried out in a human-modified landscape of Caatinga dry forest in northeast Brazil. We compiled information on woody plant assemblages inhabiting a forest chronosequence consisting of 28 forest stands of varying age: 14 regenerating (4–50-yr old) and 14 old-growth forests. Woody plant assemblages along forest regeneration had higher occurrence of specialized traits and strategies, such as pollination by vertebrates (16.6%) and dispersion by animals (47.8%). Regenerating forest stands exhibited a higher occurrence of species with specialized floral size (19%) and pollination by vertebrates (26.5%) as compared to old-growth forests. Regenerating forest stands also supported a higher (30%) functional diversity of reproductive traits than old-growth stands. Biomass, aridity, and chronic anthropogenic disturbances affected a myriad of traits/strategies: pollination by birds and animal dispersion increased with increases in biomass while total functional diversity responded negatively; pollination by Sphingids and beetles negatively responded to aridity, while very large fruit size species negatively responded to chronic anthropogenic disturbances. Both forest biomass and aridity correlated negatively with reproductive trait diversity. The Caatinga dry forest regeneration suggests high resilience, no directional/predictable changes (although regenerating vs. old-growth differ in some traits) and aridity imposed to the successional mosaics as a driver. This pattern probably emerges conditioned by the reproductive traits posed by the plant species able to thrive in such stressful environments, including those species that persist through the entire regeneration process, i.e., the regeneration process is marked by high complexity and low predictive power. Although rapid recovery represents good news, the plant-assemblage reproductive profile imposed by disturbance-adapted species may drastically alter resource availability for a myriad of plant attendants (e.g., pollinators and thief) and, consequently, the complex/diverse plant-animal interactions typical of dry forests. This working hypothesis has both theoretical and applied implications in the context of dry forest regeneration in human-modified landscapes.

An anchovy ecosystem indicator of marine predator foraging and reproduction.

Forage fishes are key energy conduits that transfer primary and secondary productivity to higher trophic levels. As novel environmental conditions caused by climate change alter ecosystems and predator-prey dynamics, there is a critical need to understand how forage fish control bottom-up forcing of food web dynamics. In the northeast Pacific, northern anchovy (Engraulis mordax) is an important forage species with high interannual variability in population size that subsequently impacts the foraging and reproductive ecology of marine predators. Anchovy habitat suitability from a species distribution model (SDM) was assessed as an indicator of the diet, distribution and reproduction of four predator species. Across 22 years (1998-2019), this anchovy ecosystem indicator (AEI) was significantly positively correlated with diet composition of all species and the distribution of common murres (Uria aalge), Brandt's cormorants (Phalacrocorax penicillatus) and California sea lions (Zalophus californianus), but not rhinoceros auklets (Cerorhinca monocerata). The capacity for the AEI to explain variability in predator reproduction varied by species but was strongest with cormorants and sea lions. The AEI demonstrates the utility of forage SDMs in creating ecosystem indicators to guide ecosystem-based management.

Evidence of abnormal scalar timing property in alexithymia.

Evidence suggests that incidental modulation of affective states affects the ability to keep track of time. Alexithymia represents an ideal condition to further address the emotion-time processing link, as it refers to a trait characterized by a deficit of affective processing. 31 healthy participants completed an online version of the TAS-20 scale, which measures alexithymia, and a time reproduction task of visual stimuli related to positive (i.e., happiness) and negative (i.e., anger) facial expressions. Results documented a positive correlation between TAS-20 score and the variability in reproducing sub-second durations of the anger expression stimuli We also found an overestimation of sub-second durations of non-affective expressions in borderline/alexithymic participants. Finally, in line with the literature, we confirmed the overall tendency to overestimate the duration of anger expression stimuli. These findings, which can be interpreted in terms of abnormal scalar timing property in alexithymia, expand previous investigations linking this personality trait with abnormal processing of negative emotions. The evidence that alexithymia predicts the reproduction variability of sub-second durations of negative affective stimuli corroborates previous neuroimaging studies documenting cerebellar deficits in these individuals.

The influence of conditions in Lake Superior and the Bois Brule River, Wisconsin on returns of migratory rainbow trout

Rainbow trout were introduced to Lake Superior in the late 1800’s and exhibit a potamodromous life history and exhibit high variability in reproductive success. We examined reproductive variability in the Bois Brule River, WI (Lake Superior), through analyses of returns of wild first spawning (hereafter “maiden” returning) adults. We used classification and regression tree analyses to identify in-stream and in-lake (western Lake Superior) sources of variability and to identify the environment (stream or lake) that was most influential to the returns to each location. Among in-stream influences, high discharge rates in the spring period (March – May) during a pre-smolt’s first stream year were the strongest source of variability and were negatively correlated with returns. High discharge during the fall period from September to November in the pre-smolt first stream year was also negatively correlated with numbers of maiden returning steelhead from that year class. When variables associated with Lake Superior were considered, maiden returns were positively correlated with higher lake surface temperatures in Lake Superior. Returns were negatively correlated with the abundance of adult rainbow smelt and bloater suggesting a possible competitive interaction among those species. Finally, we also observed a conditional (minor) positive effect of age-0 smelt abundance indicating the importance of this prey for juveniles in colder years in western Lake Superior. Taken together, our findings indicate that both streamand lake conditionsin their first lake year are important sources of variability and point to spates in the spring and fall as initialcontrolling variables.

The role of sea ice in the distribution, habitat use, and phenology of eastern North Pacific gray whales

A strong negative relationship has been observed between calf production of eastern North Pacific gray whales Eschrichtius robustus and Pacific Arctic sea ice area with the exception of 2 potentially anomalous periods: 2013-2014 and 2017-2019. Sea ice may play a role in reproductive variability by blocking gray whale access to foraging hotspot habitats. We tested this ‘sea ice exclusion’ hypothesis using complementary aerial survey and moored hydrophone data sets that together provide broad spatial and temporal coverage. A negative relationship (generalized additive model, p < 0.001) was found between gray whale aerial counts and ice concentration in the northeast Chukchi Sea (2008-2019), with a nonlinear increase in negative slope above 45-55% concentration. A comparison of distribution patterns between years further indicated the absence of sightings within northern Bering and northeastern Chukchi foraging hotspots during years with delayed ice break-up versus years with an early-to-average ice retreat. Passive acoustic recordings likewise showed the near-absence of detections during periods of dense ice cover and indicated a strongly positive relationship between ice break-up dates and the timing of acoustic detection onset. Together, these analyses point to the important role of sea ice in gray whale distribution, habitat use, and phenology. However, a relatively consistent 10-15 d lag observed between ice break-up and acoustic onset dates suggested that the mechanism(s) underpinning gray whale interactions with sea ice may be more complex than simply ‘sea ice exclusion.’ Here, we propose ‘prey quality timing’ as a potential alternative hypothesis that warrants further investigation.

Anatomy and histological evaluation of the reproductive system of marine calanoid copepod Centropages furcatus from a mass culture perspective

Marine finfish culture is a priority area for future human food security. Although many marine finfish species have been domesticated and successful breeding and spawning in captive conditions are achieved, the larval rearing from hatchling to fingerling stage to the desired level is yet to be achieved. One of the essential impediment factors in this process is the availability of suitable live feed for the finfish larvae. The traditional live feeds (Artemia nauplii and rotifers) must be more adequate in size spectrum and nutritive value to many marine finfish larvae. In nature, copepod nauplii, copepodite stages, and adults constitute the preferred food of marine finfish larvae. Copepod size spectrum, nutritive value and swimming movements make them ideal live prey items for fish larvae. Nevertheless, the high-density culture of copepods is challenging due to their sexual reproduction and high species-specific variability. Further, adequate knowledge of the candidate species' food and feeding habits, reproductive biology, and life cycle strategies should exist. With this objective present study describes the female and male reproductive system, oogenesis and spermatogenesis, egg production, and reproductive potential of a candidate calanoid copepod species, Centropages furcatus. This basic information will help develop mass culture protocol for this species.

Factors influencing the timing and frequency of litters in captive fennec foxes (Vulpes zerda)

Maintaining reproductive seasonality can be vital to the fitness of wild animals. Certain species, however, may display aseasonal reproduction and may produce multiple yearly litters when maintained in captivity. Wild fennec foxes (Vulpes zerda), for example, produce a single litter in March or April although their reproductive behaviors are reportedly variable in captivity. Here, we used the fennec fox studbook to extract traits related to reproductive variability in 220 captive-born litters. The captive litters in our dataset were born during every month of the year and nearly half (47%) were born outside of the expected months. The production of multiple litters in a single year was common, where 67% of the litters represented a second annual litter for a given dam. We detected several traits related to multi-litter years, including the dam’s ability to habituate to the birth location, the dam’s age, the dam’s number of previous litters, and the dam’s birth month. Although producing multiple litters within a year has been anecdotally associated with the loss of a previous litter, we did not detect a relationship between multi-litter years and the survivorship of previous litters. These findings suggest that captive populations of fennec foxes may experience a destabilization of their reproductive patterns, which may culminate in the production of multiple litters outside of the typical window of reproduction. Fennec foxes are a common captive species bred in zoos worldwide. Developing a greater understanding of their reproduction can allow for more successful captive management, which may improve future reproductive successes.

Genetically derived effective population size estimates of herpetofaunal species should be used with caution

AbstractEffective population size (NE) is an important parameter for conservation of threatened wildlife. Current estimators of NE that use genetic data, however, may not be equally useful among taxa that are difficult to sample, such as cryptic, iteroparous, long‐lived herpetofauna. To better understand the use of NE estimators for such taxa, we generated and compared contemporary NE estimates to evaluate methods, repeatability, and the effects of sample size at management scales and functional genetic population scales for the eastern box turtle (Terrapene carolina carolina) and eastern hellbender (Cryptobranchus alleganiensis alleganiensis). We chose these 2 species for their differing dispersal abilities, reproductive variances, and natural history traits, which may be useful in exploring the weaknesses and strengths of NE estimators. We collected species‐specific microsatellite genotypes from 2,712 individuals (1,133 hellbenders, 1,579 box turtles) in 2007–2011 from across their ranges in the eastern United States. We then grouped samples into range‐wide genetic clusters, state‐level management populations, and small regional management populations (e.g., state parks). We also randomly subsampled (rarefied) from these 3 categories to explore the effects of reduced sampling effort on NE estimates. We used 4 popular estimators: NEESTIMATOR (linkage disequilibrium and heterozygote excess methods), LDNE, and ONESAMP. Over half of all attempts at estimating NE failed to give complete estimates (those that included only non‐negative and non‐infinity values). There was variation between species but also among estimators. Repeated sampling of genetic populations resulted in inconsistent results, and estimates generally increased with sample size. The linkage disequilibrium version of NEESTIMATOR returned the largest percentage (95.8%) of complete estimates. Program ONESAMP obtained complete estimates for 62.0% of attempts and generally provided the numerically largest estimates. Only 19.4% of LDNE estimates and none of the heterozygote excess version of NEESTIMATOR estimates were complete. Generally, NE estimates were higher for box turtles, perhaps because of their greater dispersal capabilities, historically larger populations, and longer lifespans. Our results suggest that estimates of NE for long‐lived herpetofauna species vary according to species, sample size, genetic cluster, management population, and estimator used. Therefore, we recommend managers of populations of long‐lived herpetofaunal species use NE estimators with caution, and consider results from multiple methods before incorporating NE into management practice.

Polygamy and purifying selection in birds

Good genes theories of sexual selection predict that polygamy will be associated with more efficient removal of deleterious alleles (purifying selection), due to the alignment of sexual selection with natural selection. On the other hand, runaway selection theories expect no such alignment of natural and sexual selection, and may instead predict less efficient purifying selection in polygamous species due to higher reproductive variance. In an analysis of polymorphism data extracted from 150-bird genome assemblies, we show that polygamous species carry significantly fewer nonsynonymous polymorphisms, relative to synonymous polymorphisms, than monogamous bird species (p = .0005). We also show that this effect is independent of effective population size, consistent with the alignment of natural selection with sexual selection and "good genes" theories of sexual selection. Further analyses found no impact of polygamy on genetic diversity, while polygamy in females (polyandry) had a marginally significant impact (p = .045). We also recapitulate previous findings that smaller body mass and greater geographic range size are associated with more efficient purifying selection, more intense GC-biased gene conversion, and greater genetic diversity.

Parasitism shapes selection by drastically reducing host fitness and increasing host fitness variation.

Determining the effects of parasites on host reproduction is key to understanding how parasites affect the underpinnings of selection on hosts. Although infection is expected to be costly, reducing mean fitness, infection could also increase variation in fitness costs among hosts, both of which determine the potential for selection on hosts. To test these ideas, we used a phylogenetically informed meta-analysis of 118 studies to examine how changes in the mean and variance in the outcome of reproduction differed between parasitized and non-parasitized hosts. We found that parasites had severe negative effects on mean fitness, with parasitized hosts suffering reductions in fecundity, viability and mating success. Parasite infection also increased variance in reproduction, particularly fecundity and offspring viability. Surprisingly, parasites had similar effects on viability when either the male or female was parasitized. These results not only provide the first synthetic, comparative, and quantitative summary of the strong deleterious effects of parasites on host reproductive fitness, but also reveal a consistent role for parasites in shaping the opportunity for selection.