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Advertisement-call preferences in diploid-tetraploid treefrogs (Hyla chrysoscelis and Hyla versicolor): implications for mate choice and the evolution of communication systems.

Signals used for mate choice and receiver preferences are often assumed to coevolve in a lock-step fashion. However, sender-receiver coevolution can also be nonparallel: even if species differences in signals are mainly quantitative, females of some closely related species have qualitatively different preferences and underlying mechanisms. Two-alternative playback experiments using synthetic calls that differed in fine-scale temporal properties identified the receiver criteria in females of the treefrog Hyla chrysoscelis for comparison with female criteria in a cryptic tetraploid species (H. versicolor); detailed preference functions were also generated for both species based on natural patterns of variation in temporal properties. The species were similar in three respects: (1) pulses of constant frequency were as attractive as the frequency-modulated pulses typical of conspecific calls; (2) changes in preferences with temperature paralleled temperature-dependent changes in male calls; and (3) preference functions were unimodal, with weakly defined peaks estimated at values slightly higher than the estimated means in conspecific calls. There were also species differences: (1) preference function slopes were steeper in H. chrysoscelis than in H. versicolor; (2) preferences were more intensity independent in H. chrysoscelis than in H. versicolor; (3) a synergistic effect of differences in pulse rate and shape on preference strength occurred in H. versicolor but not in H. chrysoscelis; and (4) a preference for the pulse shape typical of conspecific calls was expressed at the species-typical pulse duration in H. versicolor but not in H. chrysoscelis. However, females of H. chrysoscelis did express a preference based on pulse shape when tested with longer-than-average pulses, suggesting a hypothesis that could account for some examples of nonparallel coevolution. Namely, preferences can be hidden or revealed depending on the direction of quantitative change in a signal property relative to the threshold for resolving differences in that property. The results of the experiments reported here also predict patterns of mate choice within and between contemporary populations. First, intraspecific mate choice in both species is expected to be strongly influenced by variation in temperature among calling males. Second, simultaneous differences in pulse rate and pulse shape are required for effective species discrimination by females of H. versicolor but not by females of H. chrysoscelis. Third, there is greater potential for sexual selection within populations and for discrimination against calls produced by males in other geographically remote populations in H. chrysoscelis than in H. versicolor.

Acoustic Stimulation↗

Signal-receiver interplay in the communication of male condition by Asian elephants.

Signal design and meaning are dependent on the condition of the sender and receiver as well as the response of the receiver. This study examined (1) whether female Asian elephants, Elephas maximus, can distinguish between a conspecific male in musth and nonmusth states using urinary signals, (2) how the oestrous condition of the female affects discrimination, and (3) correlation of female responses with the testosterone level of the male. Musth is a rut-like state displayed by healthy adult male elephants. Males in musth dominate nonmusth males and may be preferred by females as mates. Urine was collected from two captive male Asian elephants during nonmusth periods and from one of these males during times of musth. Samples of musth and nonmusth urine and control liquids were placed in an elephant enclosure weekly for 16 weeks, the length of a female oestrous cycle. Primary response behaviours were approach and four trunk-tip motions, namely sniff, check, place and flehmen. Musth urine consistently elicited greater responses than nonmusth and control samples. Females were more responsive during their follicular (sexually receptive) than luteal (unreceptive) stages of oestrus. Furthermore, females appeared to be sensitive to the degree of musth as responses increased with rising serum testosterone levels of the male donor. Chemical signals from males are a likely source of honest signals related to status and reproductive condition. Female elephants appear capable of detecting differences in a male based upon urinary chemosignals. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Floral evolution and pollinator mate choice in a sexually deceptive orchid.

Sexually deceptive orchids mimic sex pheromones and appearance of female insects to attract males, which pollinate the flowers in an attempted mating. This study examines the effects of pollinator mate choice on orchid floral evolution using the Thynnine wasp Neozeleboria cryptoides (Smith) (Hymenoptera: Tiphiidae), which pollinates the sexually deceptive orchid Chiloglottis trapeziformis Fitzg. (i) When male wasps were given the choice between two female dummies of different sizes and identical amount of synthetic pheromone, they preferentially attempted to copulate with medium-sized dummies over small dummies. (ii) When given the choice between two dummies of identical size but different amounts of pheromone, males preferred the larger amount of pheromone. Larger amounts of pheromone generally attracted more males than smaller amounts. (iii) Orchid flower labella, which mimic a female body, were significantly longer and broader than female wasp bodies, and the flowers also produced on average 10 times more 'pheromone' than females. The evolution and maintenance of these exaggerated mating signals is likely to be mediated by the male pollinator behaviour demonstrated here. (iv) When five dummies were offered simultaneously in a 10 cm circular array, males rarely attempted copulation on more than one dummy during a single visit. This behaviour may foster the evolution or maintenance of clonality in C. trapeziformis, as it will minimize pollen exchange within clones.

Animals↗

Neural networks predict response biases of female túngara frogs.

Artificial neural networks have become useful tools for probing the origins of perceptual biases in the absence of explicit information on underlying neuronal substrates. Preceding studies have shown that neural networks selected to recognize or discriminate simple patterns may possess emergent biases toward pattern size of symmetry--preferences often exhibited by real females--and have investigated how these biases shape signal evolution. We asked whether simple neural networks could evolve to respond to an actual mate recognition signal, the call of the túngara frog, Physalaemus pustulosus. We found that not only were networks capable of recognizing the call of the túngara frog, but that they made remarkably accurate quantitative predictions about how well females generalized to many novel calls, and that these predictions were stable over several architectures. The data suggest that the degree to which P. pustulosus females respond to a call may often be an incidental by-product of a sensory system selected simply for species recognition.

Animal Communication↗

Dopamine D2 receptor-mediated regulation of partner preferences in female prairie voles (Microtus ochrogaster): a mechanism for pair bonding?

This study examined the role of dopamine (DA) in partner preference (PP) formation in female prairie voles (Microtus ochrogaster). The nonspecific DA antagonist haloperidol blocked mating-induced PP, whereas the nonspecific DA agonist apomorphine induced PP without mating. The D2 antagonist eticlopride, but not the D1 antagonist SCH23390, blocked PP, whereas the D2 agonist quinpirole, but not the D1 agonist SKF38393, induced PP without mating. Injections of eticlopride before or immediately after mating, but not 24 hr after mating, impaired PP, indicating that DA's effects were not due to an interference with mating or sensory recognition. Finally, intracerebroventricular injections of eticlopride diminished PP. Together, these data suggest that mating-induced PP requires activation of D2 receptors and that social experience may activate dopaminergic pathways, with enduring effects on behavior.

2,3,4,5-Tetrahydro-7,8-dihydroxy-1-phenyl-1H-3-ben↗

Chemical signals in the reproduction of Asian (Elephas maximus) and African (Loxodonta africana) elephants.

Asian (Elephas maximus) and African (Loxodonta africana) elephants exhibit polygynous mating that involves female choice of mates and male-male competition for access to females. Chemical signals mediate intersexual and intrasexual interactions associated with reproduction. The need for reliable and honest signals is accentuated by the markedly different social structure of adult males and females. Adult female elephants live in matriarchal herds consisting of a dominant female and several generations of offspring. Adult males are solitary or travel with other males except during breeding periods. Because females have a long 16-week oestrous cycle with a brief 1-week receptive period and a 4-5 year interval between births, a sexually active female is a limited resource. Asian elephant females advertise a forthcoming ovulation by releasing (Z)-7-dodecen-1-yl acetate in their urine during the preovulatory period. African elephants probably produce a sex pheromone as well. Females regularly contact the ano-genital region of other females and show heightened chemosensory responsiveness to urine during the follicular phase. The physiological impacts of this ability to detect reproductive condition (e.g. possible synchronizing or suppressing of oestrus) are uncertain. Males experience an annual period of heightened aggressiveness and highly elevated testosterone concentrations known as musth. Males secrete fluid copiously from their temporal gland and dribble strongly odoriferous urine during musth. Females appear to prefer musth males as mates, and captive Asian females exhibit greater chemosensory responses to urine from males in musth than not. Males in musth are competitively dominant to all other males, even those larger than themselves. Nonmusth males avoid males in musth, and captive Asian bulls show greater interest in musth than nonmusth urine. In captivity subordinate Asian females back away from musth secretions, and females with calves sometimes display protective behaviour. Clearly, chemical signals play an important role in communication by elephants between and within the sexes. Further work is needed to identify more of these chemical messengers and to understand their complete function in mediating reproductive interactions in the elephant social system.

Animals↗

Chimpanzees are indifferent to the welfare of unrelated group members.

Humans are an unusually prosocial species-we vote, give blood, recycle, give tithes and punish violators of social norms. Experimental evidence indicates that people willingly incur costs to help strangers in anonymous one-shot interactions, and that altruistic behaviour is motivated, at least in part, by empathy and concern for the welfare of others (hereafter referred to as other-regarding preferences). In contrast, cooperative behaviour in non-human primates is mainly limited to kin and reciprocating partners, and is virtually never extended to unfamiliar individuals. Here we present experimental tests of the existence of other-regarding preferences in non-human primates, and show that chimpanzees (Pan troglodytes) do not take advantage of opportunities to deliver benefits to familiar individuals at no material cost to themselves, suggesting that chimpanzee behaviour is not motivated by other-regarding preferences. Chimpanzees are among the primates most likely to demonstrate prosocial behaviours. They participate in a variety of collective activities, including territorial patrols, coalitionary aggression, cooperative hunting, food sharing and joint mate guarding. Consolation of victims of aggression and anecdotal accounts of solicitous treatment of injured individuals suggest that chimpanzees may feel empathy. Chimpanzees sometimes reject exchanges in which they receive less valuable rewards than others, which may be one element of a 'sense of fairness', but there is no evidence that they are averse to interactions in which they benefit more than others.

Aging↗

Crickets with extravagant mating songs compensate for predation risk with extra caution.

Modern models for the evolution of conspicuous male mating displays assume that males with conspicuous displays must bear the cost of enhanced predation risk. However, if males can compensate behaviourally for their increased conspicuousness by acting more cautiously towards predators, they may be able to lower this cost. In the field cricket Gryllus integer, males call to attract females, and differ in their durations of uninterrupted trilling (calling-bout lengths). Differences among males in calling-bout lengths are heritable, and females prefer males with longer calling bouts. In this study, males with longer, more conspicuous songs behaved more cautiously than males with shorter songs on two different tests of predator avoidance. They took longer to emerge from a safe shelter within a novel, potentially dangerous environment, and they ceased calling for a longer time when their calls were interrupted by a predator cue. Thus, these males appear to compensate behaviourally for their more conspicuous mating displays. Additionally, latencies to emerge from a shelter in the novel environment were consistent over time for both individual males from the field and males that had been reared in the laboratory, indicating that the differences in latency among males may be heritable.

Animals↗

Dual functions of a melanin-based ornament in the common yellowthroat.

Melanin-based ornaments often function as signals in male-male competition, whereas carotenoid-based ornaments appear to be important in female mate choice. This difference in function is thought to occur because carotenoid pigments are more costly to produce than melanins and are thus more reliable indicators of male quality. We examined the role of melanin- and carotenoid-based ornaments in male-male competition and female choice in the common yellowthroat Geothlypis trichas, a sexually dichromatic passerine. Males display a black facial mask produced by melanin pigmentation and a bright yellow bib (throat, breast and belly) produced by carotenoid pigmentation. In controlled aviary experiments, mask size was the best predictor of both male-male competition and female mate choice, and, therefore, mask size may be regarded as an ornament of dual function. These dual functions may help to maintain the reliability of mask size as an indicator of male quality, despite the potentially low cost of production. The size of the bib was unrelated to male-male competition or female choice, but there was a tendency for females to prefer males with more colourful bibs. We propose that the black mask is important in competition for territories with other males and for attracting females. Our results highlight the need for more studies of the mechanisms of sexual selection in species with ornaments composed of different pigment types.

Animal Communication↗

Host-plant-derived variation in ultraviolet wing patterns influences mate selection by male butterflies.

We report on the first case in which sequestered secondary plant compounds determine an insect's external appearance in the ultraviolet spectrum and thereby influence visually mediated mate choice. Larvae of the common blue butterfly Polyommatus icarus specifically sequester flavonoids in different amounts and types, depending on the part or species of food plant. During late pupal development the majority of ultraviolet-absorbing flavonoids are deposited in the wing scales. The flavonoid content of the larval diet thereby determines ultraviolet wing patterns. In laboratory and field experiments, male butterflies clearly preferred flavonoid-rich, ultraviolet-absorbing female dummies. This preference is mediated visually by the ultraviolet pattern of the wings. Food-plant parts and species vary in value as a food source, so ultraviolet wing patterns may signal mate quality and are not a species-specific characteristic. We discuss the use of principal component analysis in analysing spectral data in the context of visual communication. We propose the alternative application of confidence intervals of averaged spectra as a novel straightforward statistical method for comparing groups of spectra in a manner that is independent of assumptions about the visual system of the receiver. In addition, they can be used to give confidence intervals to derived measures of colour such as quantum catch by photoreceptors.

Animals↗

The logic of Ménage à Trois.

Recent studies of socially monogamous species have shown that in many cases females do not copulate exclusively with their pair mates, but are also receptive to other males. The explanation usually given for unfaithful female behavior is that most females are unable to bond with a male they would prefer as genetic father to their offspring. To secure male assistance the female therefore pairs with an available male but also copulates with males of supposedly higher genetic quality. Here we offer an alternative evolutionary explanation for female infidelity, which does not rely upon this 'Good Genes hypothesis of female choice. We show with a simple model that in an evolutionary game between three players, a male, a female and a male lover, solutions exist in which the female can secure more assistance from her mate by being receptive to other males. We conclude that female sexuality can have a decisive role in regulating social behaviour, in which the fertile female is the driving force.

Animals↗

Individual recognition of female hamsters by males: role of chemical cues and of the olfactory and vomeronasal systems.

Sexually satiated male hamsters preferred to investigate and to mount an anesthetized, estrous, novel female over a similarly presented female with which the male had become satiated (the Coolidge effect); likewise, such males preferred a novel female recently mated with another male over the familiar female but showed no preference between fresh and mated novel females. Thus the Coolidge effect is at least partly dependent on discrimination of a new female by chemical cues. Another experiment indicated that transfer of a male's own scent during mating is not involved in discrimination between familiar and novel females. Flank gland secretion of females were sufficient for individual discrimination by males, whereas head region scents and vaginal secretions were not sufficient. The presence of female's flank glands was not, however, necessary for such discrimination. Lesions of or removal of the vomeronasal organ did not disrupt the preferences of sexually satiated males for a novel female, but elimination of main olfactory system function by ZnSO4 treatment of the olfactory mucosa did abolish such preferences. Thus olfactory cues are sufficient for individual discrimination of novel females by sexually satiated male hamsters, and such recognition leads to increased sexual arousal. These processes are mediated by the main olfactory system but not the vomeronasal accessory-olfactory system.

Animals↗

Female preferences in a fish genus without female mate choice.

The evolution and the adaptive logic (if any) of female mate choice are subjects of lively debate. Whereas most researchers believe that females have evolved to recognize signs of male 'quality' (the ability to provide females or their offspring with direct or indirect genetic or material benefits), there is intriguing evidence that males can evolve to appeal to pre-existing female preferences. Evidence for these pre-existing biases is often ambiguous because phylogenetic reconstructions have usually failed to establish conclusively whether the female preference or the favored male traits evolved first. This potential difficulty is minimal in the mosquitofish genus Gambusia, none of whose 45 species appears to have a female-choice mating system in the wild, and none of which shows the male behavioral and morphological traits that are characteristic of female choice. Nevertheless, in an experimental situation in the laboratory, female Gambusia holbrooki readily chose between models of males and demonstrated significant and reliable preferences for a variety of exaggerated male traits that are not seen in their species or their genus. Other morphological alterations were not preferred. The latent willingness of females to choose traits in a genus without such traits and without evident female choice in the wild is remarkable and may indicate a pre-existing bias in females that is ready to drive male evolution, should the social system or the ecological variables that control it change.

Animals↗

Effect of intrauterine position and social density on age of first reproduction in wild-type female house mice (Mus musculus).

In laboratory strains of mice, reproductive maturation is influenced by both the fetal and the peripubertal social environments experienced by females. Intrauterine position (IUP) for female fetuses is identified by the number of adjacent males; where 0M have zero, 1M have one, and 2M have two male neighbors. We sought to confirm, using wild-type female house mice (Mus musculus), the previous finding in the CF-1 strain of laboratory mouse that males prefer 0M over 2M females as mates. We recorded the age at first reproduction for females of known IUP housed either individually with a male or in a group of 6 females and a male. In neither housing condition were there significant differences in age of first reproduction among females of different IUPs, but 0M females that were group housed had perforate vaginas at an earlier age than other group-housed females. Mean (+/- SE) age at first reproduction was 58.1 (+/- 3.3) for paired and 126.6 (+/- 5.1) for group-housed females. The reproductive suppression observed in the grouped females was probably due to the interaction of inhibitory urinary chemosignals, low body weights, and female dominance hierarchies.

Animals↗

Sexual selection and the fitness consequences of male body size in the seed beetle Stator limbatus

We examined sexual selection on male body size in a laboratory population of the seed beetle, Stator limbatus, and the fitness consequences to females of mating with larger males. Large males produced larger ejaculates than small males. Both males and females lost body weight as a consequence of breeding, and large males lost more weight than small males. The amount of weight lost by males correlated as highly with female fecundity as did the amount of weight lost by females. Similarly, male and female body weight correlated equally highly with female fecundity. These results indicate that males make substantial contributions to female fecundity, probably through nutrients transferred in their ejaculate. As a consequence, fecundity selection should favour large body size in both males and females. We found no preference for large males when virgin females were presented with only one male, but when presented with two males simultaneously, females were more likely to mate with the larger male. This result is consistent with relative female choice or male-male competition, although no indications of male-male competition were observed. Females that mated with small males re-mated sooner than females that first mated with large males. Females that first mated with a non-virgin male were also more likely to re-mate than females that first mated with a virgin male, suggesting that females re-mate to obtain additional sperm or nutrients and not just as a form of mate choice. In addition to the possible benefits from mate choice and male-male competition, large males gain a mating advantage through reduced sperm competition. This large male advantage, combined with fecundity selection on males as well as females, may account for males being larger than females in this species.Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Female barn owls (Tyto alba) advertise good genes.

The good genes hypothesis of sexual selection postulates that ornamentation signals superior genetic quality to potential mates. Support for this hypothesis comes from studies on male ornamentation only, while it remains to be shown that female ornamentation may signal genetic quality as well. Female barn owls (Tyto alba) display more black spots on their plumage than males. The expression of this plumage trait has a genetic basis and it has been suggested that males prefer to mate with females displaying more black spots. Given the role of parasites in the evolution of sexually selected traits and of the immune system in parasite resistance, we hypothesize that the extent of female plumage 'spottiness' reflects immunological defence. We assessed the genetic variation in specific antibody production against a non-pathogenic antigen among cross-fostered nestlings and studied its covariation with the plumage spottiness of genetic parents. The magnitude of the antibody response was positively correlated with the plumage spottiness of the genetic mother but not of the genetic father. Our study thereby provides the first experimental support, to our knowledge, for the hypothesis that female ornamentation signals genetic quality.

Animals↗

Bulls fail to show preference for estrous females in serving capacity tests.

Eighteen sexually experienced Hereford bulls were exposed individually to a pair of restrained hormone-induced estrous and nonestrous females to determine mating preferences in the context of serving capacity testing. No evidence was obtained to support the hypothesis that restrained estrous females are more attractive to bulls than restrained nonestrous females. Estrous and nonestrous females received similar numbers of mount intentions, mount attempts, mounts without ejaculations and ejaculations during the 20-min tests. Time spent with estrous and nonestrous females and latencies to leave initially chosen estrous and nonestrous females to investigate the alternative female also were similar. Bulls interacted more with the females that they contacted first, regardless of estrous condition. A female position effect was noted. We conclude that restrained nonestrous females are as effective as restrained estrous females in assessing the serving capacity of Hereford bulls.

Animals↗

Effects of formalin on bacterial growth in mink feed, feed consumption and reproductive performance of adult mink, and growth of mink kits.

Feed that is typically used on commercial mink ranches is an ideal environment for bacterial growth because of the raw animal by-products used as ingredients. Recently, formaldehyde was approved for use as an antimicrobial agent in poultry feed. Experiments in our laboratory were carried out to investigate the effects of incorporating different concentrations of formalin into the feed of mink on the growth of gram-negative and gram-positive bacteria. Feed containing 0, 550 or 1100 ppm formalin was kept refrigerated for up to 7 d and the number of colony forming units of gram-negative and gram-positive bacteria derived from the feed was determined each day. Colony forming units in the formalin-treated feed were significantly fewer than colony forming units in untreated feed. In the second trial, feed containing the same concentrations of formalin was maintained at 30 C for 24 h and cultured bacterial colonies were counted at 0, 12 or 24 h of feed incubation. Both concentrations of formalin were effective in significantly reducing the number of colony forming units. A feed consumption trial determined if mink (Mustela vison) preferred formalin-treated feed to non-treated feed kept refrigerated for up to 7 d. Consumption of feed treated with 1100 ppm formalin was significantly lower than consumption of the non-treated feed on d 1, 2, 4 and 5, but body weight was not affected. A long-term feeding trial determined the effects of formalin on mink reproduction, early growth of offspring and quality of fur. Mink were fed formalin at concentrations of 0, 550 or 1100 ppm for approximately 140 d beginning 1 mo prior to mating until kits were weaned at 6 w of age. Mating success was not affected by consumption of formalin-treated diets, but kit survival at birth was adversely affected in mink consuming 1100 ppm formalin. Hemoglobin concentration, hematocrit, mean corpuscular volume, and mean corpuscular hemoglobin were significantly decreased in 6-w-old kits, but there were no significant differences in any of these parameters between the kits exposed to 0 and 550 ppm formalin. In a second phase, some kits and their dams were continued on their respective dietary treatments from weaning through pelting (approximately 220 and 320 d, respectively). At pelting, hematocrits and hemoglobin concentrations for the kits fed 1100 ppm formalin were significantly less compared to the control and 550 ppm formalin groups. There were no significant differences in body weights among female kits or adult female mink. The body weights of male kits in the 1100 ppm formalin group became significantly less than the body weights of male kits in the control and 550 ppm formalin groups as the trial progressed. The quality of fur was highest for mink in the control group and lowest for mink in the 1100 ppm formalin group. While dietary 1100 and 550 ppm formalin were effective in suppressing bacterial growth in the feed of mink, the deleterious effects of 1100 ppm formalin on kit survival, hematologic parameters, body weight, and quality of fur preclude formalin use at this concentration.

Animal Feed↗