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AB - Abstract Mate choice is generally regarded as an independent event, but a growing body of evidence indicates that it can be influenced by social information provided by conspecifics. This is known as non-independent mate choice. Individuals use information gathered by observing interactions between conspecifics to copy or not copy the mate choice of these conspecifics. In this review, we examine the factors that affect non-independent mate choice and mate choice copying and how it is influenced by social and environmental information that is available to the subject or focal individual. Specifically, we discuss how non-independent mate choice and whether individuals copy the choices of conspecifics can be influenced by factors such as habitat and differences in ecology, mating system and parental care. We focus on the social information provided to the focal animal, the model and the audience. Nearly all studies of non-independent mate choice and mate copying have focused on individuals in species that use visual cues as the source of social information. Nevertheless, we highlight studies that indicate that individuals in some species may use chemical cues and signals as sources of social information that may affect non-independent mate choice and mate copying. audience effect context, focal individuals, mate choice copying, non-independent mate choice, social information INTRODUCTION Independent mate choice Classical theories surrounding mate choice stipulate that an individual will choose a particular mate based on features of its genotype and phenotype that are attractive to that individual and that these traits and preferences for them have been selected for over generations (Darwin, 1859, 1871; Fisher, 1930; Zahavi, 1975; Kodric-Brown & Brown, 1984). Regardless of the mechanism(s) involved in selecting a mate, these theories imply that the chooser is making a selection independently, i.e. without any further information provided by conspecifics or possibly heterospecifics. The net result is that individuals are more likely to choose conspecifics that they perceive as being higher quality mates than lower quality mates, and by doing so gain indirect and direct fitness benefits (Andersson, 1994). Thus, a central requirement of these classical theories of mate selection is that an individual’s ability to identify and to discern variation among suitors is contingent upon that individual’s ability to recognize and then choose high-quality opposite-sex conspecifics (Andersson & Simmons, 2006). Typically, preferences for such traits have a strong genetic component and are usually independent of factors that may influence them such as the behaviour and choices of conspecifics (Bakker & Pomiankowski, 1995; Mead & Arnold, 2004; Andersson & Simmons, 2006). For example, asymmetries in the size of two potential mates can be used by opposite-sex conspecifics to choose mates. Male Trinidadian guppies (Poecilia reticulata), littorinid gastropods (Littorina subrotundata), pot-bellied seahorses (Hippocampus abdominalis) and sand lizards (Lacerta agilis) prefer larger females over smaller females, independent of those of conspecifics (Olsson, 1993; Zahradnik et al., 2008; Mattle & Wilson, 2009; Godin & Auld, 2013). The explanation given for such a preference is that males can maximize their fitness by mating with larger females because larger females are generally more fecund. Such preference by males for larger females will cause selection pressures on females to become larger and presumably more fecund (Andersson, 1994; Godin & Auld, 2013). Similarly, differences in the coloration of conspecifics can be used as a mechanism for distinguishing quality among potential mates. Female house finches (Carpodacus mexicanus) preferred more colourful males to less colourful males independently of the male’s age or social status (Hill, 1990). Female guppies preferred male guppies with brighter pigmentations over male guppies with duller pigmentations (Kodric-Brown, 1985). The coloration of male finches and guppies is dependent on the amount of carotenoids in their diet (Goodwin, 1950; Hill et al., 2002), which is an indicator of that male’s ability to secure high-quality forage (Kodric-Brown, 1985). Thus, the brightness of the coloration of males in these species may be considered an honest signal of male quality (Kodric-Brown, 1985), and preferences for such males should be independent of preferences displayed by conspecifics for other potential mates. Other traits such as sound production and chemical cues that indicate phenotype and genotype can be used to discern among potential mates. Female water pipits (Anthus spinoletta) (Rehsteiner et al., 1998), European starlings (Sturnus vulgaris) (Eens et al., 1991), wolf spiders (Hygrolycosa rubrofasciata) (Kotiaho et al., 1996) and field crickets (Gryllus lineaticeps) (Wagner & Reiser, 2000) have all been found to select mates based on certain aspects of the males’ songs/calls such as duration and complexity. In addition, meadow voles (Microtus pennsylvanicus) preferred the scent marks of adult, opposite-sex conspecifics that have higher titres of free gonadal steroids relative to those that have lower titres (Ferkin et al., 2017). Male and female meadow voles with higher gonadal sex steroid titres are more likely than those with lower titres of such hormones to be successful sires and dams (Ferkin et al., 2017). House mice (Mus musculus) prefer mates with a genetically dissimilar major histocompatibility complex to those with similar major histocompatibility complex (Eklund, 1997). Presumably, this mating preference functions to provide ‘good genes’ for an individual’s offspring because the major histocompatibility complex regulates immunological self/non-self recognition (Penn & Potts, 1999), and histocompatibility complex heterozygosity enhances reproductive success (Thoss et al., 2011). Collectively, these studies have shown that preferences for oppoiste-sex conspecifics favors those with better genes and phenotypes and preference for such higher quality individuals are typically independent of those for mate choices demonstrated by conspecifics. Non-independent mate choice The signals and cues used for mate choice are often directed at a specific target(s), but can also be detected by non-target individuals (Dugatkin, 1992; Danchin et al., 2004; Valone, 2007). These signals and cues may provide public information to rival conspecifics. The unintended (non-target) receiver may then respond to these signals and cues, modifying their behaviour towards the sender or the targeted receiver based on the information obtained (Danchin et al., 2004; Witte et al., 2015; Ferkin & Ferkin, 2017). Thus, behavioural interactions between opposite-sex conspecifics are also a source of social information to individuals (Danchin et al., 2004; Valone, 2007). This social information gained by observing conspecifics may be used by an individual to influence its mate choice, if that individual has no personal information about those conspecifics (Ferkin, 2018, 2019a, b). Thus, the net effect is that individuals may use personal information, social information or both to facilitate mate choice, with personal information providing more accurate and reliable information than would social information alone. A growing body of evidence shows that social information may also affect mate choice, resulting in non-independent mate choice (Dugatkin, 1992; Pruett-Jones, 1992; Witte et al., 2015). Some of the first empirical evidence for non-independent mate choice, specifically mate choice copying, began to surface during the early 1990s with work by Dugatkin & Godin (1992) studying guppies from Trinidad. In their study, female guppies were placed in the centre of an aquarium with two aquaria containing males of similar length on opposite ends juxtaposed to the central aquarium. The female guppy was able to swim freely in the aquarium in which she was placed. The amount of time she spent near each male was compared and the preferred male was the one with whom she spent the most time near. Once a preference had been established, a model female was placed behind a Plexiglas divider next to the originally non-preferred male; the preferred male received no model. The focal female could then observe both her originally preferred and non-preferred male. After the observation, the model female was removed and the focal female was given the opportunity to choose between the two males again. Dugatkin & Godin (1992) found that female guppies switched their preference towards the initially non-preferred male by spending more time near him. Thus, the association of the non-preferred male with a model female was sufficient to sway the focal female’s preference from the originally preferred male and towards the ‘choice’ of the other female, in this case the previously non-preferred male. More importantly, female guppies used the social information gathered from observing an association between conspecifics to switch their preference for a male. The change in preference by the focal female was induced by the social information she gained by observing the behavioural interaction of the model female and the male she originally did not prefer. Thus, the females are not making an independent mate choice. Rather, they are using social information to make a non-independent mate choice (Dugatkin, 1992; Pruett-Jones, 1992; Witte et al., 2015). In this review, we want to move beyond whether a species engages in mate choice copying and delve into the specific factors that affect non-independent mate choice and mate choice copying. Specifically, we discuss the factors that may impact the mate choice decisions of focal individuals. These potential factors include the effects of features of the habitat and local environment, the mating and social system of the participants, the type of social information provided by the model that is responded to by the focal animal, the context in which the social information is sent and received, and whether an audience is present. We discuss how these factors can be affected by the social information they can provide to the focal individual. In addition, we use a growing literature to highlight gaps in our knowledge about the factors behind non-independent mate choice and mate choice copying, and where possible to delineate areas of consensus and lack of consensus. As we explored the literature on non-human animals, we discovered that the vast majority of studies of non-independent mate choice have been conducted on fish and birds, and to a lesser degree, on insects and mammals. We chose not to include human studies in our review. An excellent, detailed and current review on non-independent mate choice and mate copying in humans has been written by Gouda-Vossos et al. (2018). In addition, most of the studies that have examined non-independent mate choice have focused on species that use visual cues and signals as sources of social information. Some of these biases are by necessity reflected in our review. However, we discuss the results of a slowly growing number of studies, indicating that olfactory and chemical cues and signals provide social information that can affect non-independent mate choice and mate copying in a variety of non-human animals. FACTORS AFFECTING NON-INDEPENDENT MATE CHOICE AND DECISIONS TO MATE COPY Role of habitat and local environment Features of the habitat and local environment in which a species resides shapes their reproductive behaviours, including mate choice (Endler, 1995; Caro et al., 2009). The differences in features of the habitat and local environment may be the factor behind conflicting reports on mate choice copying in guppies. Studies of guppies from Australia (Brooks, 1999), South Africa (Brooks, 1996) and a stock purchased from a pet store (Lafleur et al., 1997) failed to find mate choice copying using similar methods to those of Dugatkin & Godin (1992). We do not know the origins of the Australia and South Africa guppy populations and whether those habitat and/or environmental constraints placed on populations in Australia and South Africa have negated mate copying as a suitable behaviour. Perhaps, the cost of expressing non-independent mate choice and mate copying no longer provides benefits. Thus, it is difficult to tell how the environment has shaped the mate choice behaviours from this population. It would be interesting to know if captive guppies from Trinidad, Australia and South Africa engage in non-independent mate choice and in mate copying if placed in identical conditions. Doing so would test Brooks’ (1996, 1999) hypothesis that there is a strong genetic component for mate copying in guppies and that the local habitat affects the occurrence of non-independent mate choice. Predation risk is another aspect of the environment that may affect independent mate choice and mate choice copying. Female guppies from parts of the Quaré River, Trinidad, where there is a high predation risk, were less likely to independently choose brightly coloured males when a predator pike cichlid was present (Crenicichla alta) (Godin & Briggs, 1996). In contrast, female guppies from a river with low predation risk, the Paria River, were not affected by the presence of a predator pike cichlid and continued to independently prefer the brighter male (Godin & Briggs, 1996). Briggs et al. (1996) found that female guppies from the high-predation-risk population from the Quaré River were not affected by the presence of a pike cichlid and engaged in mate choice copying. Briggs et al. (1996) suggested that female guppies engage in mate choice copying even when predation risk is high because of the reduced time and cost of assessing potential mates that they would otherwise incur. This hypothesis supposes that for the guppies in Trinidad the benefits of copying outweigh the costs of not copying. This is consistent with aspects of sexual selection and mate choice theory (Endler, 1995) that under increasing predation risk females may reduce their exposure to predators by searching less extensively for a mate (Hedrick & Dill, 1993). In this case, a female guppy may reduce her predation risk by relying on the choice of conspecifics for mates (Briggs et al., 1996). For many species during the breeding season, competition for breeding or nesting sites can place pressure on individuals to reproduce (Slagsvold & Viljugrein, 1999). It could potentially affect the engagement of mate choice copying. Female pied flycatchers (Ficedula hypoleuca) did not show a preference for a male that associated with a model female, and instead the focal females made their choice independently based on the courting behaviour of the males (Slagsvold & Viljugrein, 1999). One conclusion drawn was that limited nest-site availability would accentuate the need for a female pied flycatcher to find a mate quickly, potentially leaving little time for her to survey the choices of other females (Slagsvold & Viljugrein, 1999). Thus, mate choice copying would not be beneficial for female pied flycatchers because the costs of gathering social information for choosing a mate may lead to loss of nest-site acquisition. One aspect not discussed by Slagsvold & Viljugrein (1999) is how the density of nesting sites and/or the population could affect the mate choice decisions of pied flycatchers. Both of these variables can create vastly different scenarios of the availability of social information, which then could influence the potential use of said social information. For example, if nest-sites are not localized and are spread out, an individual’s cost of obtaining social information may be high in terms of energy expended and risk of predation. On the other hand, if nest-sites are limited to specific areas or patches, the cost for an individual to gathering social information may decrease. It would be interesting to see if the variables related to the spatial distribution of nest-sites and population density could affect whether an individual will rely on social information for mate choice decisions. In high-density populations, individuals may gain more social information. However, the social information may not be as reliable as it would be in low-density populations. In low-density populations, the cost of providing unreliable social information may be high because individuals are more likely to have subsequent interactions with the same conspecifics than they would have in high-density populations. In a recent study, Dagaeff et al. (2016) discovered that atmospheric pressure influenced mate copying in female Drosophila melanogaster. Female fruit flies were more likely to mate copy under high and increasing air pressure relative to low and decreasing air pressure. The authors suggested that the increasing air pressure would probably signal to the D. melanogaster that favourable weather conditions were approaching. It would be of interest to see if changes in air pressure and other features of local weather influence the willingness of individuals of other species to engage in non-independent mate choice. We have presented results suggesting that features of the habitat and local environment will probably affect whether individuals will use social information provided by conspecifics and heterospecifics to affect mate choice decisions. However, the number of such studies is limited. More studies should examine the role of habitat and local conditions and what specific information the focal individual uses to engage in non-independent mate choice and whether they copy the mate choice of conspecifics. By doing so, we can create models that predict under what conditions and features of the local habitat we would expect to see mate choice copying. Role of mating system Mating system can affect the type and frequency of social information available to individualsm which may influence their mate choice decisions. Vakirtzis (2011) compiled results of studies for several species and categorized them as to whether mate choice copying was found, not found or inconclusive. Vakirtzis (2011) also included information about the mating system of the species. The vast majority of studies of mate choice copying were conducted on species with a promiscuous mating system; fewer studies were conducted on species that were considered to be polygynous and monogamous. Vakirtzis (2011) identified that among the promiscuous species studied, most showed mate choice copying. The sand goby (Pomatoschistus minutus) (Forsgren et al., 1996) and common goby (Pomatoschistus microps) (Reynolds & Jones, 1999) did not show mate copying. Similarly, Vakirtzis (2011) found conflicting reports of mate choice copying in two monogamous species. Pied flycatchers were reported to not mate copy (Slagsvold & Viljugrein, 1999). However, zebra finches (Taeniopygia guttata) were found to mate copy in one study (Drullion & Dubois, 2008) but not in another (Doucet et al., 2004). In addition, Vakirtzis (2011) reported that only two species with polygynous mating systems have been studied and both the house mouse (Mus musculus) (Kavaliers et al., 2006) and a marine isopod (Paracerceis sculpta) (Shuster & Wade, 1991) showed evidence of mate choice copying. At present, we can find no consensus among studies as to whether mating systems are a good predictor of whether a species engages in non-independent mate choice. This concern is highlighted by the fact that many studies examined the likelihood that non-independent mate choice would occur in only one sex (Forsgren et al., 1996). Perhaps, males and females within the same species may differ in whether they will engage in non-independent mate choice and whether the factors affecting such a decision are the same. It is also possible that the mode of fertilization, external or internal, affects the propensity for mate choice copying or non-independent mate choice to occur. However, data are limited to support either of these potential hypotheses. Role of parental care Another aspect related to the effects of mating system on non-independent mate choice decisions is the degree to which males and females provide parental care. Generally, females are considered the choosier sex due to their generally higher initial reproductive investment (Trivers, 1972), which could explain why non-independent mate choice is tested more often in female but not male birds and fish. However, in many species of fish, males but not females provide parental care (Breder & Rosen, 1966). Thus, Frommen et al. (2009) posited that mate copying should be more common in species that display parental care and in the sex that provides more parental care. Presumably, mate choice copying will occur more frequently in the sex that provided the greater reproductive expenditure. Moran et al. (2013) studied banded darter (Etheostoma zonale) and fantail darters (E. flabellare) and found that the male fantail darter mate copied whereas the female fantail darter did not. In this species, male but not female fantail darters provide parental care for the developing eggs (Page & Swofford, 1984). In contrast, male and female banded darters engage in non-independent mate choice and mate copy but neither sex provides parental care (Moran et al. 2013). Widemo (2005) studied mate choice in male and female deep-snouted pipefish (Syngnathus typhle), a live bearing species, and discovered that male but not female pipefish mate copied. Note also that male but not female pipefish provide parental care. Currently, the hypothesis postulated by Frommen et al. (2009) has gained mixed support. Clearly, it is too soon to tell if the sex that provides more parental care will be more likely than the other sex to engage in non-independent mate choice and whether non-independent mate choice is prevalent in males and females in species in which neither sex provides parental care. Role of social information and its reliability A model individual provides three forms of information to the focal individual: the model’s individual phenotype and genotype, the behaviour of a model towards an opposite-sex conspecific, and the behaviour of the opposite-sex conspecific towards the model. These pieces of information can affect the use of social information by an individual for their mate selection(s). However, whether the focal individual will respond in a particular manner depends on whether it is able to determine if the cues provide information and if the information provided is reliable (Wheeler et al. 2011; Ferkin, 2018, 2019a, b). Features of the phenotype and genotype of a potential rival or model provides social information that can influence the mate choice of individuals (Kodric-Brown, 1985; Penn & Potts, 1999; Thoss et al., 2011). Male guppies chose a female guppy that was adjacent to two drab males over a female that was adjacent to two colourful males (Gasparini et al., 2013). The focal male guppies appear to be able to gauge their competitive ability in conjunction with the likelihood of being selected when choosing a mate. A focal male guppy that is colourful compared to a drab-coloured male conspecific is more likely to be viewed as being of high quality and chosen as a mate by a female guppy. However, the same focal male may have a lower likelihood of being selected by that female as a mate if she is surrounded by other colourful male conspecifics. In this case, the focal male may switch his preference and attempt to mate with the less-preferred or lower quality female because she would be less likely to reject him. Switching by the focal male may represent a trade-off between the cost of mating with a lower quality female and the benefit of having a mating opportunity with a lower risk of sperm competition than if he attempted to mate with the preferred female. Nordell & Valone (1998) noted that females are more likely to copy when it is hard to discriminate between high- and low-value mates, which may influence mate choice copying. Japanese medakas (Oryzias lattipes) provide an example in which the type and reliability of social information provided can affect the likelihood of inducing mate choice copying. Grant & Green (1996) allowed focal female medakas to observe a model female spawn with a male that the focal female did not prefer initially. After watching the spawning, the focal female later showed a preference for that male, indicating that she switched her choice of mates and copied the choice of another female. In contrast, Howard et al. (1998) reported that when focal female Japanese medakas observed a model female with an extruding clutch of eggs, presumably giving the impression to the focal female of having already spawned with the non-preferred male, the focal females did not switch their preference to the male they initially did not prefer; the focal female medakas did not exhibit mate-copying. The extruded clutch of eggs from the model female may not have been a reliable indicator that that female had recently mated with the male. By contrast, in the Grant & Green (1996) study, the observation of spawning was a reliable signal and was sufficient for the focal female to copy the mate choice. Similarly, mate choice in male Atlantic mollies (Poecilia mexicana) was influenced by social information obtained by observing a model male interact with a female conspecific. Focal male Atlantic mollies were found to prefer an originally non-preferred female when a model male was in close proximity but could not interact with a female, whereas that preference did not occur when the model male interacted with the female (Bierbach et al., 2011b). Bierbach et al. (2011b) suggested that the male’s strong willingness to mate choice copy when the model was not physically interacting with the female could be attributed to the focal male’s perception of that female as being a lower risk for sperm competition. The type and reliability of the social information presented to focal individuals can also affect mate choice decisions in birds. Höglund et al. (1990) conducted field observations of the lekking behaviour of black grouse (Tetrao tetrix) and suggested that the skewness seen in the mating success of males was based on mate choice copying by female black grouse. Höglund et al. (1995) later tested this hypothesis by placing decoys within the territory of a male black grouse before the female grouse arrived at the lek. In their experiment, Höglund et al. (1995) used black grouse decoys that were placed on the ground, or placed above the ground, or duck (heterospecifics) decoys on the ground. Male black grouse that encountered black grouse decoys on the ground later received more attention by female grouse compared to the attention gained by male black grouse that encountered black grouse decoys above the ground or that encountered duck decoys (Höglund et al., 1995). Thus, the location as well as the type of information conveyed by the social interaction between the male and the decoy affected the amount of time that female black grouse would attend to them. In black grouse, the presumption of copulation with another female was sufficient for focal females to show a preference for a male that they considered as having copulated with the model female relative to a male that they considered as not having copulated with the model female. Similarly, both curly-winged and wild-type female D. melanogaster preferred a curly-winged male after observing that male copulate (Nöbel et al., 2018). Typically, female wild-type and curly-winged D. melanogaster will not show a preference for curly-winged mutant males over wild-type males (Nöbel et al., 2018). A question that remains is why would females of both phenotypes show a preference for a mutant that can result in decreased fitness? It may be that observing the act of copulation provides reliable information that the focal female D. melanogaster uses for their subsequent choice. This speculation is in line with the results from the Grant & Green (1996) study of Japanese meddaka and the Höglund et al. (1995) study of black grouse. Encounters with a heterospecific or observations of interactions involving a heterospecific could also affect the mate choice of the focal individual. Typically, animals live in areas with multiple species present. Thus, different species can interact and provide cues to focal individuals. Amazon mollies (Poecilia formosa) are a gynogenetic hybrid of Atlantic mollies and sailfin mollies that need copulatory behaviours from a male of either species to trigger embryogenesis (Hubbs & Hubbs, 1932; Balsano et al., 1989; Schlupp et al., 2002). Female sailfin mollies changed their preference to an initially non-preferred male after viewing the preferred male with a female Amazon molly (Hill & Ryan, 2006). Hill & Ryan (2006) suggested that the switched preference may be due avoidance of the preferred male instead of the focal individual developing a preference for the non-preferred male. Thus, we do not know if female sailfin mollies avoided preferred males that had associated with a heterospecific Amazon molly. In contrast, female mate choice in Drosophila serrata was independent of whether that male associated with either conspecific females or female Drosophila birchii. Female D. serrata chose males randomly with no signs of mate choice copying (Auld et al., 2009). Clearly, species differences exist in how focal individuals that respond to conspecifics that are associated with heterospecifics. Thus, the question remains as to whether focal individuals in some species cannot or choose not to place any value on cues provided by heterospecifics. In this way, the cues from heterospecifics would not provide social information to focal individuals. Alternatively, focal individuals in some species cannot determine if social information involving heterospecifics is reliable or warrants their attention. Nearly all studies conducted on non-independent mate choice and mate choice copying have focused on the focal individual observing a conspecific having a positive interaction with the model. However, non-independent mate choice and mate choice copying may also be affected by a focal individual’s observation of an encounter that a model individual had which resulted in a negative interaction with a conspecific. Observing a negative interaction may be sufficient to induce a focal individual to not select or choose to associate with an opposite-sex conspecific. For example, female D. melanogaster rejected a male they observed being rejected by a model female (Mery et al., 2009) and female sailfin mollies will reject males that were observed with a female attempting to escape from them (Witte & Ueding, 2003); a similar phenomenon of rejection has been observed in humans (Hill & Buss, 2008). We need to test the hypothesis that social information that reflects negative interactions between the model and an opposite-sex conspecific will reduce the likelihood of non-independent mate choice. We may need to consider whether individuals differ in their threshold to interpreting particular interactions between conspecifics as being negative, neutral or positive. Assigning such a valence to an interaction may depend on the context of the observation, the nature of the social information provided, the condition of the participants and the physiological/emotional state of the focal individual. Several factors can affect the reliability of information provided to the focal individual. For example, the size and age of the model can have profound effects on mate choice decisions. Focal female guppies copied when the model was larger but not smaller than the focal female (Vukomanovic & Rodd, 2007). Female guppies also copied if the model female was older but not younger than the focal female (Dugatkin & Godin, 1993). The probable reason for the copying of larger or older models over smaller or younger models is that size and age provide a greater assurance of conveying accurate information to the focal individual. A larger or older model is probably more experienced in making mate choice decisions and may have a better ability to accurately assess potential mates. This in turn would mean that the reliability of the information gleaned from those individuals will more probably be accurate and reliable. For the focal individual that would mean the benefits of using this information increases while the costs decrease. The reliability of social information provided to the focal individual may also depend on the amount of time that it can view social interactions involving the model. Female sailfin mollies prefer large males over small males, independent of an observed 10-min association the model may have with a small male (Witte & Ryan, 1998; Witte & Noltemeier, 2002). However, if the focal female sailfin molly was allowed to observe the model with a small male for 20 min she copied the choice of the model and showed a preference for the small male over the large male (Witte & Noltemeier, 2002). Clearly, the amount of time that the sailfin molly observed the small male and model affected her mate choice. The question is why? It is possible that longer viewing times lead to an increased perception of reliability of the social information the model is providing to the focal individual. Being able to view conspecific interactions for a longer period of time increases the strength of the cue being provided; in this case viewing a female associating with a male for a longer period of time increases the strength of her perceived preference for that male. The interval of time between the observation of the model and a conspecific and the test for mate choice also affected whether an individual mate copied. Female D. melanogaster, male Japanese quail (Coturnix japonica) and male guppies no longer preferred an opposite sex-conspecific directly after viewing it copulate but did prefer this individual after 24 h (Loyau et al., 2012), 48 h (White & Galef, 2000) and 24 h (Makowicz et al., 2010), respectively. For focal female individuals, the avoidance of a male after initially observing copulation would potentially prevent copulation with a male that is sperm-depleted. For focal male individuals, the avoidance of a female after initially observing copulation would prevent undesired sperm competition with other males. The temporal changes in preference seen with D. melanogaster, Japanese quail and guppies suggest social learning and memory play a key role in affecting non-independent mate choice and mate choice copying. Unreliable information and misinformation So far, we have provided examples in which the focal individual received reliable social information by the model. However, instances may occur when the model, knowingly or not, provides inaccurate and unreliable social information to the focal individual. For example, female zebra finches appear to copy only when the social information they obtained was most likely to be reliable. In the study by Drullion & Dubois (2008), female zebra finch models were given different leg bands to create ‘reliable’ and ‘unreliable’ information. The authors found that focal female zebra finches only copied when the male and female had the same-coloured leg bands; presumably, the focal female zebra finch considered this information to be reliable. Focal female zebra finches did not mate copy if the male and female had different-coloured leg bands. In this case, information was considered unreliable by the focal females (Drullion & Dubois, 2008). In another example, Kniel et al. (2017) altered the phenotypes of male and model female zebra finches by placing red feathers on top of their heads. Focal female zebra finches no longer chose a male she initially preferred if the male had a red feather placed on his head (Kniel et al., 2017). The focal female may have interpreted the model female to be of another population or possibly another species (Kniel et al., 2017). Thus, the red feather added to the head of the male may have provided social information suggesting that the male was a heterospecific. However, we can consider that for the female zebra finches in this experiment, the cues from the ‘seemingly heterospecific male’ actually provided reliable information that would have prevented a failed mating. Collectively, these studies suggest that mate choice decisions of the focal individual can be affected by the type of information provided by the model and conspecifics interacting with the model (regardless of whether the information is reliable), and is attended to by the focal individual. Some data exist that suggest that individuals may provide a rival(s) in the audience with misinformation. In this case, the misinformation a male may provide to its rivals is that the selected female was not the preferred female. Providing misinformation may be a form of deceptive behaviour on the part of the model individual. Deception, at least for humans, occurs when a sender produces a signal that it knows to be false with the intention of a false belief in receivers (Wheeler et al., 2011). For non-human animals, deception ignores the intention of the sender. Few studies have been conducted to examine the deception hypothesis. In a study by Plath et al. (2010), the authors concluded that surface-dwelling but not cave-dwelling mollies were presenting deceptive signals. We do not know if males from the studies mentioned were intending to deceive their rivals by associating with the former non-preferred female. This speculation presumes that surface-dwelling mollies possess higher cognitive abilities or a theory of mind. Nevertheless, it is possible that for these animals the model (sender) derives a fitness benefit by signalling false information to potential rivals (e.g. Searcy & Nowicki, 2005; Wheeler et al., 2011). The fitness benefit male fish may obtain by indicating a preference for a ‘less desirable’ female in the presence of a rival may be a greater chance to attempt to mate with this female. In that way, the sender may increase his paternity assurance and reduce sperm competition when he mates with the ‘desirable’ female. A recent study that set out to test the deception hypothesis found evidence that focal male Atlantic mollies reversed their original preference for a female if an audience was present (Witte et al., 2018). They also found that the audience male showed a preference for the female that the focal male associated. Witte et al. (2018) concluded that the behaviour of the focal individual supported the deception hypothesis. In addition, these authors suggested that an individual would not alter its behaviour unless there was a large cost associated with the risk of being copied (Witte et al., 2018). A lowered risk of sperm competition for the focal individual is clearly a benefit. However, if deception is a viable tactic, its occurrence should be infrequent, dependent on population density, and provide a large benefit but a large potential cost to the actor. Thus, the actor should be more likely to target naïve and unfamiliar individuals and less likely to target experienced and familiar individuals. Reliability of the information provided by the model is important in the cost–benefits of using social information to influence mate choice decisions by the focal individual. The costs of using unreliable social information or failing to correctly interpret this information could have dramatic effects on the reproductive success of the focal individual. In addition, receiving misinformation would be costly to the focal individual as it may lead to lost mating attempts and unnecessary predation risk. Providing misinformation inadvertently or intentionally is also likely to affect mate choice decisions and ultimately reproductive success, and potentially the survival of the focal individual (Witte et al., 2018). As experience may affect how an individual responds to social information (Ferkin, 2019a, b), one could hypothesize that the age and experience of the focal individual should affect what social information is sufficient to induce it to mate copy. Presumably, older and more experienced individuals would be less susceptible to social information that may be questionable or provided by an unreliable model. In addition, studies need to be conducted that measure the costs and benefits to the model of providing reliable or unreliable social information. Perhaps, older individuals are more likely than younger individuals to provide reliable social information to older conspecifics as the latter would be unlikely to be deceived by unreliable information. Older individuals may be more likely than younger individuals to provide unreliable information to younger conspecifics. Individuals may have to learn to which individuals they can or cannot provide unreliable social information. Audience effect Individuals may use information gleaned from observing interactions among conspecifics to aid in their own decision-making (Valone, 2007; Vakirtzis, 2011; Witte et al., 2015), but in doing so, observers may affect the behaviour of the individuals being watched. If one or both individuals of a social interaction are aware they are being observed they may alter their behaviour(s); this is referred to as the audience effect (Zajonc, 1965). The audience effect can influence a wide variety of reproductive behaviours. For example, males of some bird species will be alter their extra-pair copulations and behaviours towards female conspecifics if they are being observed by their own mate (Baltz & Clark, 1997; Ung et al., 2011). Similarly, lower-ranking individuals may be more likely than higher-ranking individuals to attempt to conceal their copulations for fear of reprisals by higher-ranking conspecifics (Townsend et al., 2008; Townsend & Zuberbuhler, 2009; Overduin-de Vries et al., 2012). In this section, we discuss studies of the audience effect with regard to non-independent mate choice. Male zebra finches spent more time near an initially non-preferred female, including a female bearing a different phenotype via a red feather adornment, when a rival male was present (Dubois & Belzile, 2012; Kniel et al., 2016). A hypothesis derived by both Dubois & Belzile (2012) and Kniel et al. (2016) was that the change of behaviour was related to the chance of being rejected by a preferred mate. That is, the male was showing a preference for a female that may be less likely to reject him. This hypothesis posits an interesting speculation that the individual is able to gauge its own phenotype and compare it to a rival and then tailor its own mate choice accordingly. In doing so, a male individual that views itself as lower quality relative to a rival may choose to mate with a less desirable conspecific to avoid competing with his rival, and in do so, increase its chances of reproductive success. Future studies should examine this phenomenon by comparing how individuals behave when the condition of the rival varies in comparison with the focal individual. Aspects of information context, including amount and type of audience, of the rival conspecific may influence the behaviour of focal individuals. Male sand gazelle (Gazella marica) initially changed their mate preference by spending similar amounts of time with both females when a rival was present; however, this equivocal response was lost after the first hour the rival was present (Wronski et al., 2012). The authors suggested that the focal male’s equivocal response for the first hour may have been the result of two things. The first was that the rival had no access to the females and the second was habituation of the rival male’s presence to the focal male (Wronski et al., 2012). In another study, the type of rival, conspecific or heterospecific, affected the mate preference of Atlantic mollies. The focal male mollies spent similar amounts of time with both females if a conspecific rival was present but not if a heterospecific, a green swordtail (Xiphophorus hellerii), was present (Plath et al., 2008). Male guppies showed differences in the frequency of courtship behaviour depending on the audience males’ attractiveness rating (Órfão et al., 2019). Focal male guppies had a higher frequency of courting the female when the audience male was either small or possessed large amounts of orange coloration (Órfão et al., 2019). Interestingly, the audience, regardless of their quality, had no effect on the frequency of unsolicited mating attempts by the focal males (Órfão et al., 2019). The audience effect also alters sexual behaviours performed by the focal male. Male guppies reduced the number of thrusts and nipping/approaches towards larger females when an audience was present (Makowicz et al., 2010). Focal male sailfin mollies decreased their preference for the preferred male if a rival was present but not when no rival was present. However, focal male sailfin mollies transferred significantly more sperm to the female when the rival was present compared to when the rival was not present (Nöbel & Witte, 2013). Cave-dwelling and surface-dwelling Atlantic mollies decreased their nipping and gonopodial thrusts towards the larger female when a rival was present with the surface-dwelling mollies showing the greater decrease in these behaviours (Plath et al., 2010). Surface-dwelling mollies interacted significantly more often with the smaller female when a rival was present (Plath et al., 2010). Bierbach et al., (2011a) found that male Atlantic mollies appear to remember the activities of their rival and adjust their preferences for a female conspecific. In that study, focal male Atlantic mollies were initially allowed to view rival males either alone or with females. This allowed male mollies to assess whether a rival male was sexually inactive or sexually active. Focal male Atlantic mollies altered their preference when the rival male was viewed as being sexually active but not if it was viewed as not being sexually active (Bierbach et al., 2011a). Ziege et al. (2012) also showed that focal male Atlantic mollies will change their sexual behaviours towards females if the rival conspecific interacted with the females. Focal males reduced their nipping and gonopodial thrusting towards females that were observed interacting with a rival male. Ziege et al. (2012) suggested that the changes in behaviour by the focal males were probably driven by sperm competition and that the focal male mollies avoided female mollies that they believed to have been inseminated by a rival male. Being able to gauge the activity of rivals may be a form of assessing the risk of sperm competition (Vaughn et al., 2008; Ziege et al., 2009; Parker & Pizzari, 2010). The number of rivals in the audience is often a factor that is not considered in studies of the audience effect. Most studies use an audience of only a single rival. However, additional rivals may affect the focal individual’s response to the audience. Yet, published studies offer no consensus with regard to supporting this notion. For example, no additive audience effect was reported for male guppies when the audience was increased from one to two rival males (Auld & Godin, 2015). Focal males for both the one-male and two-male audiences decreased their association times with a preferred female but there was no difference between the two groups (Auld & Godin, 2015). They suggested that an increase in one audience member may not signify a strong enough increase in the intensity of sperm competition to affect a male’s choice of a mate. However, when looking at sexual behaviours Auld et al. (2015) found no audience effect with a one-male audience compared to a control no audience but did find that a two-male audience caused focal males to decrease in courtship behaviour and sneak copulations. The results reported by Auld et al. (2015) contradict those reported by Makowicz et al. (2010), who found differences in sexual behaviour of male guppies exposed to a rival male. Multiple rivals may have a different effect than compared with a single rival on mate preference and behaviour of focal individuals. In some cases, focal individuals exposed to a rival did not alter their behaviour towards an opposite-sex conspecific they preferred originally. Focal male Grijalva mosquitofish (Heterophallus milleri) and male eastern mosquitofish (Gambusia holbrooki) did not switch their preference when a rival male conspecific was present (Ziege et al., 2008; Callander et al., 2011; Bierbach et al., 2012). The behaviour by the focal Grijalva mosquitofish was attributed to monopolization of the female (Ziege et al., 2008; Bierbach et al., 2012). Callander et al. (2011) suggested four explanations as to why eastern mosquitofish did not change their preference for a female when a rival was present. The first was that it was not an adaptive response for the focal male in that the number of rival males associated with a female eastern mosquitofish varies widely. A second explanation was that given that female mosquitofish store sperm, multiple males may still increase their reproductive success in subsequent broods by mating with a high-quality female. A third explanation was that aspects of fitness other than the number of offspring sired may play a role, such as size of the offspring. Larger females will often produce larger offspring. Thus, guarding a larger female may ensure a male has the opportunity to sire larger offspring, which would probably increase his fitness. The last explanation was that the lack of a change in preference was due to the potential elevation of other aspects of fitness that are associated with a living in a large group such as a reduction in predation risk (Callander et al., 2011). These explanations have yet to be tested, but will give insight into the effect of an audience on non-independent mate choice. It would be interesting to determine if there is a point at which the size and make-up of the audience no longer has an effect on the mate choice of a focal individual. Non-independent mate choice by using chemical signals and scent marks Studies looking at non-independent mate choice and mate choice copying have tended to focus on animals that use vision as their primary sensory modality for communicating information to conspecifics. However, many terrestrial mammals use olfaction and chemical signals as their primary means for communicating (Thiessen & Rice, 1976; Johnston, 1983; Ferkin et al., 2017) and for providing social information to conspecifics (Ferkin, 2018, 2019a, b). Akin to visual cues, scent marks can convey a wide variety of information including the age, sex, diet, reproductive condition and dominance status of the sender (Johnston, 1983, 2003; Hurst & Beynon, 2004; Ferkin et al., 2017). Unlike visual cues, however, scent marks can be deposited by the sender without the target individual and audience present. Scent marks are individually distinct and provide unique information about the donor (Johnston, 2003). Thus, individuals can glean the individual identity of donors that mark in an area and gain social information about them if their marks overlap or are adjacent to one another (Ferkin, 2019a, b). Scent marks can potentially function as chemical bulletin boards that create opportunities for individuals to gather social information about the phenotype and genotype of the scent donors and their interactions with other scent donors (Johnston, 2003; Ferkin et al., 2017). Given that scent marks can last for hours or days in the environment (Thiessen & Rice, 1976; Hurst & Beynon, 2004), the social information provided by chemical cues may persist for a period of time that is sufficient to influence the receiver’s mate choice (Ferkin, 2018, 2019a, b). The literature contains a slowly growing body of evidence showing that individuals can have their mate choice influenced by the odours of conspecifics (Ferkin, 2018). Females of an outbred strain of laboratory mouse (CF-1) preferred the odour of a male that had previously associated with an oestrous female compared to a male that had associated with a female not in oestrous (Kavaliers et al., 2006). Female Norway rats (Rattus norvegicus) preferred a male that had recently mated, out of sight of the focal female, over a male that had not previously mated (Galef et al., 2008). In addition, male rats that had recently mated had shorter latencies to mount and ejaculate with oestrous females than previously non-mated males (Galef et al., 2008). Male meadow voles preferred the odour of a female whose odour was associated with three or five males but not with zero, one or two males (Ferkin & Ferkin, 2017). Male meadow voles also preferred the odour of a female vole in heightened sexual receptivity, a female in post-partum oestrus, compared to that of a female voles that is not in post-partum oestrus but is still capable of breeding (Ferkin & Johnston, 1995; Vaughn & Ferkin, 2011). Male meadow voles, however, preferred the odour of a female not in post-partum oestrus if the odours of five males were associated with her odour compared to that of a female that was in post-partum oestrus (Vaughn & Ferkin, 2011). Although these studies did not directly study mate choice copying, they showed that the odour preferences of male meadow voles can be influenced by social information provided by scent marks from conspecifics (Ferkin et al., 2017; Ferkin, 2019a, b), which can affect mate choice decisions (Ferkin, 2018). Thus, the results of these studies support the speculation that non-independent mate choice and mate choice copying may be influenced by the social information provided by scent marks for terrestrial mammals (Vaughn & Ferkin, 2011). Proceeding forward, studies using chemical communication should look at the different factors to examine how chemical cues are obtained and processed along with the behaviours that are performed by the individual. One factor would be changes in reproductive behaviours when rivals are present. As mentioned earlier, male sailfin mollies deposit more sperm in a female when a rival is present. Likewise, male meadow voles mating with a female conspecific increased the amount of sperm in their ejaculate when a rival(s) odour was present relative to when no rival was present (delBarco-Trillo & Ferkin, 2004, 2006; Vaughn et al., 2008). More studies are needed to see how odours influence sperm production and transfer and how animals use the social information provided by scent marks to affect the mating decisions of focal individuals (Ferkin, 2019a, b). Another factor that could affect mate decisions is the relative quality of the odour donor. For example, male and female meadow voles prefer the scent marks of an opposite-sex conspecific that was fed a protein-rich diet over that of a donor fed a protein-poor diet (Ferkin et al., 1997). This finding is consistent with the hypothesis that higher quality individuals are more likely than lower quality individuals to advertise their condition when searching for a mate (Ferkin, 2018). In addition, the scent marks deposited by a higher quality but not a lower quality individual may be viewed as providing more reliable information to a focal individual. If so, a focal individual may be more likely to copy the mate choice of model conspecifics that they consider to be of higher quality. However, we cannot rule out the possibility that the focal individual would copy the choice of a lower quality model. If a focal individual is of higher quality than a model, the focal individual may be able to out-compete the model for access to the female. This particular explanation presumes that a lower quality individual attempts to attract mates in the presence of a higher quality individual. If the lower quality individual advertises to only lower quality individuals it would probably be more beneficial for the focal individual to seek out a higher quality mate. This may also be depend on the sex of the individual, as the costs of mating with lower quality individuals should be higher for females than for males (Trivers, 1972; Andersson, 1994). Similar types of questions can also be asked with regard to the age of the individual, and its reproductive condition, parasite load, hormone milieu, etc. Interestingly, these factors may not be mutually exclusive and thus complicate our understanding of how specific social information may be used by individuals to affect their mate choice decisions and whether they use social information to facilitate non-independent mate choice. Endocrine control of non-independent mate choice Few studies have examined the mechanisms behind non-independent mate choice. Kavaliers et al. (2017) proposed that oxytocin may influence mate choice copying behaviour due to oxytocin’s involvement in social recognition and connections with reward systems in the brain. They also addressed how oxytocin has a positive relationship with sexual interest/motivation. Evidence for this has been provided with oxytocin knockout (OTKO) mice. Both male and female OTKO mice and mice that were treated with an oxytocin receptor antagonist showed impairment in their ability to use indirect social information (Kavaliers et al., 2006). Thus, oxytocin levels can affect both the focal individual and the model individual. The focal individual may be incapable of processing the social information gathered when that individual has low levels of oxytocin. By comparison, a model with low oxytocin may not act in a typical fashion, meaning that the information the focal individual is gathering is not reliable. These speculations are consistent with the findings of Kavaliers et al. (2019). They found that female house mice not infected with a pathogen do not mate copy if the male mouse was associated with a female model that was infected with a pathogen or if both the female model and the male were infected with a pathogen. The shifts in the preferences of female house mice for males were attributed to oxytocin in facilitating preferences by the focal female for the odour of a male (Kavaliers et al., 2019). A recent study has also found that both dopamine and serotonin are necessary for female mate choice copying in D. melanogaster (Monier et al., 2019). Virgin adult female flies were modified to reduce the production of both dopamine and serotonin and upon doing so this led to random mate choice. Following treatment with the precursors of dopamine or serotonin, these female flies began to engage in mate choice copying (Monier et al., 2019). Thus, the precursors of dopamine or serotonin were necessary to reinstate non-independent mate choice in female fruit flies. It would be interesting to see if these indoleamines are also involved in facilitating mate choice copying in vertebrates. We know that the sexual behaviour of vertebrates is mediated by gonadal sex steroids, testosterone and its metabolites, and protein hormones such as vasopressin and oxytocin (Bronson & Whitten, 1968; Lombardi et al., 1976; Achiraman & Archunan, 2005). We also know that oestrogens are involved in social recognition and learning, and sexual behaviours (Clipperton et al., 2008; Ervin et al., 2015; Lymer et al., 2017, 2018). It is also likely that oestrogens may play a role in mate choice copying. Therefore, we may posit the hypothesis that females with high oestrogen titres will be more likely than those with low oestrogen titres to engage in mate choice copying. Likewise, males with high androgen titres will be more likely than those with lower androgen titres to engage in mate choice copying. Given that circulating gonadal sex steroid titres also affect the attractiveness of opposite-sex conspecifics and their willingness to mate (Nelson & Kreigsfeld, 2017), model individuals with higher titres of androgens or oestrogens may be more likely than model individuals with lower titres of these hormones to show non-independent mate choice and mate copying. Although these hypotheses are highly speculative, they do provide a framework for future testing to determine if the mechanisms underlying mate choice copying are shared by many species. FINAL THOUGHTS We have reviewed studies and highlighted the factors that may affect and underlie non-independent mate choice and mate choice copying in non-human animals. We close this review by providing scaffolding for a new framework for questions regarding non-independent mate choice and mate choice copying. A major piece of this framework is that it incorporates the importance of social information, the effectiveness of sender/model/audience to use or manipulate it, and the changing trade-offs in the value, salience and importance of that information so that the focal individual can tailor its responses accordingly, presumably to maximize it benefits. Specifically, information about conspecifics and heterospecifics can be extracted from cues and signals that are observed by others. Individuals that can detect cues, determine their reliability and respond to them may use the information they provide to influence their mate choice decisions. If the signals are reliable and accurately reveal the situation, focal individuals may use this information, if they have little personal information to select mates. However, the social information provided can be influenced by several factors, including the context, features of the habitat, the genotype, phenotype, and number of participants and observers, as well as the state of the focal individual. Individuals must be able to reconcile these factors and tailor their responses accordingly. Thus, mate choice copying and non-independent mate choice may be flexible responses to social information for individuals in many species. Differences in individual- and trait-based copying might explain the relative lack of consensus among studies examining the factors that can affect non-independent mate choice. Indeed, differences in personality among focal animals may impact whether individuals will use mate choice copying and whether their willingness to do so is stable over time and different contexts. The likelihood that mate choice copying has evolved in a particular species may be a result of the fitness benefits non-independent mate choice provides when personal information about a mate was not available. In many species individuals may copy the mate choice of conspecifics, and therefore it is possible that the trade-off in fitness between non-independent mate choice and dependent mate choice is not large and may represent an adaptive alternative strategy. Our intention is to encourage research into these factors and to encourage researchers to delve deeper into the adaptive significance, evolution, development and physiological aspects of non-independent mate choice and mate choice copying. ACKNOWLEDGEMENTS We thank Karl N. Rohrer for reading an earlier version of the manuscript, and three anonymous reviewers for providing critical and helpful comments. Writing was supported by funds from the Jack H. Morris Distinguished Professorship to M.H.F. 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TI - Factors that affect non-independent mate choice
JO - Biological Journal of the Linnean Society
DO - 10.1093/biolinnean/blz112
DA - 2019-10-18
UR - https://www.deepdyve.com/lp/oxford-university-press/factors-that-affect-non-independent-mate-choice-8rFydjrv0a
SP - 1
VL - Advance Article
IS - 
DP - DeepDyve
ER -