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The sexual selection continuum.

The evolution of mate choice for genetic benefits has become the tale of two hypotheses: Fisher's 'run-away' and 'good genes', or viability indicators. These hypotheses are often pitted against each other as alternatives, with evidence that attractive males sire more viable offspring interpreted as support for good genes and with a negative or null relationship between mating success of sons and other components of fitness interpreted as favouring the Fisher process. Here, we build a general model of female choice for indirect benefits that captures the essence of both the 'Fisherian' and 'good-genes' models. All versions of our model point to a single process that favours female preference for males siring offspring of high reproductive value. Enhanced mating success and survival are therefore equally valid genetic benefits of mate choice, but their relative importance varies depending on female choice costs. The relationship between male attractiveness and survival may be positive or negative, depending on life-history trade-offs and mating skew. This relationship can change sign in response to increased costliness of choice or environmental change. Any form of female preference is subject to self-reinforcing evolution, and any relationship (or lack thereof) between male display and offspring survival is inevitably an indicator of offspring reproductive values. Costly female choice can be maintained with or without higher offspring survival.

Alleles↗

Population structure of Anisakis simplex (Nematoda) in harbor porpoises Phocoena phocoena off Denmark.

The population structure and habitat selection of Anisakis simplex in 35 harbor porpoises off Denmark are described. The nematodes were collected from the stomach and duodenal ampulla and were categorized as third-stage larvae, fourth-stage larvae, subadults, and adults. The porpoises harbored 8,043 specimens of A. simplex. The proportion of adults and subadults increased with infrapopulation size. The number of development stages across infrapopulations covaried significantly (Kendall's test of concordance). Concordance was higher in hosts with the highest intensities than in those with low and medium intensities. All stages occurred mainly in the forestomach, but this trend was stronger for the adults. Adult and subadult sex ratios did not depart significantly from 1:1. Our data suggested that recruitment and duration of each stage were the main factors accounting for infrapopulation structure. The preference of A. simplex for the forestomach conformed with previous studies, but the narrower distribution of adults relative to other stages might indicate a strategy to enhance mating opportunities. Information on sex ratios of A. simplex is scarce and contradictory. We suggest that the discrepancies might partly reflect differences in categorization criteria and statistical methods.

Animals↗

Age-related mate choice in the wandering albatross.

We studied mate choice in the wandering albatross, Diomedea exulans, using data from 32 years of banding returns in the population of the Crozet Islands. We studied mating choices in a single year, when the Crozet Islands population was male biased (8:5, males:females). Thus, we expected that females might show great flexibility of choice of partners. Because age and experience might influence mate choice, we tested the expectation that females would choose the oldest and most experienced males for pair bonding. Pair bonds usually last until one member of the pair dies (0.3% of the birds 'divorce'), so mate choice should be especially important. We found that the ages of males and females in both displaying and bonded (breeding) pairs were significantly correlated. These age-associated pairings were not a passive phenomenon, but appeared to be due to an active process of selection of mates of similar age. First-time breeders sought mates of similar age, but preferred those with the most experience. Remating, experienced birds whose mates had died did not pair with individuals of significantly similar age, but predominantly paired with other widowed birds that, on average, were also relatively old. Mate fidelity in wandering albatrosses may be due to the cost of finding and bonding with a new mate. Pair bonds, and thus breeding, took an average of 3.2 and 2.3 years to establish, for males and females, respectively. Thus, remating exerts a potential average reproductive cost of about 15% of lifetime reproductive success. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Females use self-referent cues to avoid mating with previous mates.

Females of many species mate repeatedly throughout their lives, often with many different males (polyandry). Females can secure genetic benefits by maximizing their diversity of mating partners, and might be expected, therefore, to forego matings with previous partners in favour of novel males. Indeed, a female preference for novel mating partners has been shown in several taxa, but the mechanism by which females distinguish between novel males and previous mates remains unknown. We show that female crickets (Gryllodes sigillatus) mark males with their own unique chemical signatures during mating, enabling females to recognize prior mates in subsequent encounters and to avoid remating with them. Because self-referent chemosensory cues provide females with a simple, but reliable mechanism of identifying individuals with whom they have mated without requiring any special cognitive ability, they may be a widespread means by which females across a broad range of animal mating systems maximize the genetic benefits of polyandry.

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The effect of endotoxin-induced fever on thermoregulation in the newborn rabbit.

1. Metabolic responses of endotoxin-injected newborn rabbits were measured in a closed circuit calorimeter at a constant environmental temperature within their thermoneutral range. Oxygen consumption and colonic temperatures were then measured over a range of environmental temperatures from 21.0 to 40.0 degrees C and the responses of endotoxin-injected rabbits compared with non-injected litter mates. 2. To measure their preferred thermal environment, endotoxin-injected and non-injected litter mates were allowed to settle on a thermal gradient and their colonic temperatures measured. 3. In a constant environmental temperature of 37 degrees C, rises in oxygen consumption and colonic temperature following endotoxin injection were found to be biphasic. Oxygen consumption rose from 21 ml kg-1 min-1 to a maximum 35 ml kg-1 min-1 and colonic temperature rose from 39.0 to 39.8 degrees C. 4. The maximal rate of oxygen consumption was the same in both injected and non-injected animals, 53 ml kg-1 min-1, being provoked at the same environmental temperature of 24 degrees C. Minimal rates of oxygen consumption were also similar for the two groups but in the injected animals they were achieved at an environmental temperature of 39 degrees C, 2 degrees C higher than for non-injected animals. 5. We conclude that newborn rabbits challenged with a pyrogen have both behavioural and physiological responses. The thermogenic response is consistent with a change in sensitivity to feedback information rather than a simple shift in the central thermoregulatory set point. We find no evidence to support the view that in the febrile response a higher body temperature necessitates an increase in metabolic rate, the so-called Q10 effect.

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Degree of social contact affects the emission of food calls in the common marmoset (Callithrix jacchus).

The aim of this study was to investigate whether the emission of food calls in common marmosets (Callithrix jacchus) is influenced by different social contexts. Food calls are emitted by this species only in the presence of preferred food. If these calls have any communicative function, it may be that individuals produce food calls in order to call family mates toward the food source. If this is the case, the number of calls produced should vary in accordance with the number of family mates present at the moment of the discovery of the food, i.e., the fewer family mates nearby, the more food calls are emitted. This hypothesis was tested with five pairs of common marmosets, by recording the number of food calls emitted in four experimental conditions: 1). isolation: completely isolated from the family mates; 2). visual isolation: separated by a wooden panel from the family mates; 3). visual contact: separated by a wire-mesh from the family mates; and 4). physical interaction: together with the family mates. The results show that the proportion of intervals during which food calls were produced by the pairs was significantly different in the four experimental conditions. It decreased from the isolation and visual isolation condition, through the visual contact condition, reaching the lowest value in the physical interaction condition. The variation observed in the proportion of intervals during which food calls were emitted, in relation to different social contexts, is an indication in favor of the communicative function of this vocalization.

Aging↗

Single paternity of clutches and sperm storage in the promiscuous green turtle (Chelonia mydas).

Paternity of 22 green turtle (Chelonia mydas) clutches from 13 females of the southern Great Barrier Reef breeding population was determined through microsatellite analyses at five loci, including the analysis of successive clutches for nine of the females. A large number of alleles per locus (10-40) provided probabilities of detecting multiple paternity that were quite high, particularly at all loci combined (99.9%). Although green turtles are promiscuous breeders and there was an expectation of finding extensive multiple paternity, only two clutches were multiply sired and, in these, very few eggs had been fertilized by a secondary male. The rarity of multiple paternity may reflect either a low proportion of multiple matings by females in this population, or sperm competition, possibly resulting from a first-male sperm preference. Additionally, the analysis of > 900 offspring provided data on mutations, which included 20 mutation events that were observed in 27 offspring and involved both maternal and paternal lineages. Most mutations (n = 16) occurred at a single highly variable locus and their presence emphasizes the need to use multiple loci in paternity studies.

Animals↗

Reproductive and mate choice strategies in the hermaphroditic flatworm Echinostoma caproni.

Due to the important role that mating systems play in the evolution of species, we investigate the selfing rate and mate choice in the simultaneous hermaphroditic parasite Echinostoma caproni (Trematoda). The echinostomes were maintained in two situations in mice: (1) double infections where the two individuals do or do not belong to the same geographic area isolate, and (2) triple infections where two of the three individuals originate from the same isolate and the third one originates from a different isolate. This experimental design permits analysis of intra- and interisolate selfing rates and of mate preference. We predict, in the first experiment, no difference between intra- and interisolate selfing rates. In the second experiment we expect a preferential outcrossing between individuals originating from the same isolate in order to avoid hybrid breakdown. The results obtained corroborate our predictions and emphasize the important and synergistic roles of selfing, inbreeding depression, and hybrid breakdown in the evolution of echinostome reproductive strategies.

Animals↗

Multilocus models of sympatric speciation: Bush versus Rice versus Felsenstein.

In populations of phytophagous insects that use the host plant as a rendezvous for mating, divergence in host preference could lead to sympatric speciation. Speciation requires the elimination of "generalist" genotypes, that is, those with intermediate host preference. This could occur because such genotypes have an inherent fitness disadvantage, or because preference alleles become associated with alleles that are oppositely selected on the two hosts. Although the former mechanism has been shown to be plausible, the latter mechanism has not been studied in detail. I consider a multilocus model (the "Bush model") in which one set of biallelic loci affects host preference, and a second set affects viability on the hosts once chosen. Alleles that increase viability on one host decrease viability on the other, and all loci are assumed to be unlinked. With moderately strong selection on the viability loci, preference alleles rapidly become associated with viability alleles, and the population splits into two reproductively isolated host specialist populations. The conditions for speciation to occur in this model, as measured by the strength of selection required, are somewhat more stringent than in a model in which preference and viability are controlled by the same loci (one-trait model). In contrast, the conditions are much less stringent than in a model in which speciation requires buildup of associations between viability loci and loci controlling a host-independent assortative mating trait (canonical two-trait model). Moreover, in the one-trait model, and to a lesser extent the Bush model, the strength of selection needed to initiate speciation is only slightly greater than that needed to complete it. This indicates that documenting instances of sympatric species that are reproductively isolated only by host or habitat preference would provide evidence for the plausibility of sympatric speciation in nature.

Alleles↗

Ethological isolation and mating experience in Drosophila paulistorum.

With few exceptions, previously published experiments documenting ethological isolation between Drosophila populations have used young virgin females. We have observed directly females of Drosophila paulistorum semispecies in choice experiments with both homogamic and heterogamic males. Aged females' sexual selection did not differ from that of young females. Previous heterogamic copulatory experience did not change the degree of sexual isolation; however, females with homogamic copulatory experience showed a significantly higher preference for homogamic males.

Aging↗

Females prefer leading males: relative call timing and sexual selection in katydid choruses.

Females of several acoustic insects and anurans have recently been shown to orient preferentially to the leading of two identical male calls presented in close succession. We studied this phenomenon via two-choice phonotaxis trials in the neotropical katydid Neoconocephalus spiza and found that females avoided male calls occurring during an interval beginning immediately after the onset of a leading call. This avoidance occurred whether or not the following call was overlapped by the leader, indicating that the mechanism of preference for leading calls may represent a psychoacoustic precedence effect rather than simple call masking. We also found that females preferred leading calls as long as the amplitude of the following call was not more than 2 dB higher than the leader. Under certain circumstances, preferences for leading calls remained when females were presented with a sequence of four calls. Thus, preferences for leading calls probably influence a male's attractiveness and mating success in complex natural choruses: females are expected to orient towards males producing leading calls regardless of call length (and the probability of overlapping a following call), when the leading call is up to 1.5 times as distant as a follower, and when succeeded by multiple followers. We suggest that female preferences for leading male calls represent sensory biases that originated in contexts unrelated to sexual selection. None the less they currently play an important role in sexual selection and influence individual male signalling behaviour, and thus chorus structure. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

The shape of female mating preferences.

The "shape" of a female mating preference is the relationship between a male trait and the probability of acceptance as a mating partner. The shape of preferences is important in many models of sexual selection, mate recognition, communication, and speciation, yet it has rarely been measured precisely. Here I examine preference shape for male calling song in a bushcricket (katydid). Preferences change dramatically between races of a species, from strongly directional to broadly stabilizing (but with a net directional effect). Preference shape generally matches the distribution of the male trait. This is compatible with a coevolutionary model of signal-preference evolution, although it does not rule out an alternative model, sensory exploitation. Preference shapes are shown to be genetic in origin.

Animals↗

The influence of a male's dominance status on female choice in Syrian hamsters.

This study evaluated the importance of dominance status on mate selection in Syrian hamsters. In one experiment, sexually receptive females were allowed to choose between tethered males which differed in status. The choice was consistently in favor of the dominant male. The female spent more time in lordosis in the presence of the dominant male. The dominant also obtained a significantly greater number of intromissions. A second experiment investigated whether prior familiarization was essential to permit a female to express mating preferences in a situation where she was free to interact with three males. Again, the dominants were most often preferred and obtained greater sexual access to the female. Prior familiarization or extensive contact with the males was not necessary to support the female's selection of the dominant as a mating partner. The choice occurred quickly, generally within 5 min after contacting the males. Although females did mate with the subordinates, this typically occurred late in the tests. The significance of these data with respect to mate choice and probable paternity effects are discussed.

Animals↗

Reproduction in wild gorillas and some comparisons with chimpanzees.

Information was collected over a period of almost 12 years on the gorillas of the Virunga Volcanoes region of Rwanda and Zaire, most of it collected since September 1972. Comparisons were made with the Gombe Stream chimpanzee population (values in parentheses). Gorilla females matured at about 8 years (compared with 9-10 years for the chimpanzee) and first bred at 10-11 years (11-12 years). Males started to breed later, possibly at 15 years (about 13 years). Oestrous periods of female gorillas lasted for about 2 days (10 days) and oestrous cycles for about 30 days (36 days). Gestation in the gorilla lasts 255 days (228 days). Intervals between surviving offspring in the Virunga study groups was about 4 1/2 (5 1/2 years) but in the whole Virunga population was nearer 8 years. Lactational amenorrhoea lasted about 2 1/2 years (3 1/2 years). In gorillas and chimpanzees there were about 3 cycles to conception after parturition and females probably produce in their lifetime about 3 offspring that survive to adulthood. A successful male's productivity is greater. Mortality of immature gorillas was about 40% (50%). Initiation of courtship is generally by the female in gorillas but the male in chimpanzees. Copulation in gorillas lasts for about 1 1/2 min (7 sec) and occurs at a rate of once every 3 h when a female is in oestrus (once in 2 h). Interference in copulation is more common in the chimpanzee than the gorilla, but competition between individual males is more intense in gorilla populations. Females of both species can clearly exercise their preferences for particular males. The observed differences between the species in courtship and mating behaviour can be related to differences in the number of males available to and competing for oestrous females: in the loose multi-male chimpanzee community there is more advantage to males in initiating copulation and mating frequently and efficiently, and to females in advertising oestrus, than in the relatively stable one-male mating system of the gorilla.

Animals↗

Species and populations of the Anopheles gambiae complex in Cameroon with special emphasis on chromosomal and molecular forms of Anopheles gambiae s.s.

We studied the geographical distribution of species, chromosomal, and molecular forms of the Anopheles gambiae Giles (Diptera: Culicidae) complex in 23 sites in Cameroon, Central Africa. Almost all the specimens collected in the four northern-most arid sites were Anopheles arabiensis. Anopheles melas was found in a rural locality surrounded by mangrove swamps, on the Atlantic Coast. In total, 1,525 An. gambiae s.s. females were identified down to their molecular form, and inversion polymorphisms on polytene chromosomes were scored from 186 half-gravid females. The Forest chromosomal form, with standard arrangements almost fixed on both arms of chromosome-2, was the only one observed in the southern, more humid localities. Karyotypes typical of Savanna and Mopti were recorded northwards, in the humid savannas of the Adamawa Province. The molecular forms M and S were widespread throughout Cameroon, and assort independently from the chromosomal forms. S-form populations were characterized by karyotypes typical of Forest and Savanna chromosomal forms, and M-form populations were characterized by karyotypes typical of Forest, Savanna, and Mopti. No M/S hybrid patterns were detected, although M and S mosquitoes were sympatric in 15 sites, providing further evidence for positive assortative mating within molecular forms. The observed ecogeographical distribution of M and S was peculiar: the ecological parameters involved in this distribution still need to be clarified as well as the possible role of competitive exclusion between chromosomally homosequential molecular forms. No difference was observed in host preference or in Plasmodium falciparum infection rates between sympatric M and S populations.

Animals↗

A spatial genetic structure and effects of relatedness on mate choice in a wild bird population.

Inbreeding depression, as commonly found in natural populations, should favour the evolution of inbreeding avoidance mechanisms. If natal dispersal, the first and probably most effective mechanism, does not lead to a complete separation of males and females from a common origin, a small-scale genetic population structure may result and other mechanisms to avoid inbreeding may exist. We studied the genetic population structure and individual mating patterns in blue tits (Parus caeruleus). The population showed a local genetic structure in two out of four years: genetic relatedness between individuals (estimated from microsatellite markers) decreased with distance. This pattern was mainly caused by immigrants to the study area; these, if paired with fellow immigrants, were more related than expected by chance. Since blue tits did not avoid inbreeding with their social partner, we examined if individuals preferred less related partners at later stages of the mate choice process. We found no evidence that females or males avoided inbreeding through extra-pair copulations or through mate desertion and postbreeding dispersal. Although the small-scale genetic population structure suggests that blue tits could use a simple rule of thumb to select less related mates, females did not generally prefer more distantly breeding extra-pair partners. However, the proportion of young fathered by an extra-pair male in mixed paternity broods depended on the genetic relatedness with the female. This suggests that there is a fertilization bias towards less related copulation partners and that blue tits are able to reduce the costs of inbreeding through a postcopulatory process.

Animals↗

Contrasting mutual sexual selection on homologous signal traits in Drosophila serrata.

The nature of male mating preferences, and how they differ from female mating preferences in species with conventional sex roles, has received little attention in sexual selection studies. We estimated the form and strength of sexual selection as a consequence of male and female mating preferences in a laboratory-based population of Drosophila serrata. The differences between sexual selection on male and female signal traits (cuticular hydrocarbons [CHCs]) were evaluated within a formal framework of linear and nonlinear selection gradients. Females tended to exert linear sexual selection on male CHCs, whereas males preferred intermediate female CHC phenotypes leading to convex (stabilizing) selection gradients. Possible mechanisms determining the nonlinear nature of sexual selection on female CHCs are proposed.

Animals↗

Cytoplasmic incompatibility and mating preference in Colombian Drosophila pseudoobscura.

MACRAE and ANDERSON observed a large frequency change of mitochondrial DNA (mtDNA) haplotypes in a population initiated with two allopatric strains of Drosophila pseudoobscura, BogER from Colombia and AH162 from California. They concluded that mtDNA haplotypes in D. pseudoobscura are not always selectively neutral. NIGRO and PROUT suggested, however, that a maternally transmitted incompatibility system, similar to the one they observed in two strains of D. simulans from Italy, could account for the observed mtDNA frequency changes. SINGH and HALE postulated that a mating preference between the strains BogER and AH162 in MACRAF and ANDERSON's experiment, in the form of negative assortative mating, could also account for the mtDNA frequency changes. We report two experiments designed to test the hypotheses: that a maternally transmitted cytoplasmic incompatibility system exists between D. pseudoobscura strains BogER and AH162; and, that BogER females mate preferentially with AH162 males. Our results do not support either hypothesis.

Animals↗