Bourgeois Males Research Articles

The Evolution of Social Behaviour

The Evolution of Social Behaviour

Coevolution of female fidelity and male help in populations with alternative reproductive tactics.

In socially monogamous species, pair-bonded males often continue to provide care to all offspring in their nests despite some degree of paternity loss due to female extra-pair copulation. Previous theoretical models suggested that females can use their within-pair offspring as ‘hostages' to blackmail their social mates, so that they continue to provide care to the brood at low levels of cuckoldry. These models, however, rely on the assumption of sufficiently accurate male detection of cuckoldry and the reduction of parental effort in case of suspicion. Therefore, they cannot explain the abundant cases where cuckolded males continue to provide extensive care to the brood. Here we use an analytical population genetics model and an individual-based simulation model to explore the coevolution of female fidelity and male help in populations with two genetically determined alternative reproductive tactics (ARTs): sneakers that achieve paternity solely via extra-pair copulations and bourgeois that form a mating pair and spend some efforts in brood care. We show that when the efficiency of mate guarding is intermediate, the bourgeois males can evolve to ‘specialize' in providing care by spending more than 90% of time in helping their females while guarding them as much as possible, despite frequent cuckoldry by the sneakers. We also show that when sneakers have tactic-specific adaptations and thus are more competitive than the bourgeois in gaining extra-pair fertilizations, the frequency of sneakers and the degrees of female fidelity and male help can fluctuate in evolutionary cycles. Our theoretical predictions highlight the need for further empirical tests in species with ARTs.

Daily energy expenditure of males following alternative reproductive tactics: Solitary roamers spend more energy than group-living males

In many species, males follow alternative reproductive tactics (ARTs), where one tactic (called bourgeois) has much higher reproductive success than alternative tactics followed by males with lower competitive ability. The extent to which ARTs differ in energetic costs is unknown, but it is important to understand the fitness payoffs of ARTs. We studied male African striped mice (Rhabdomys pumilio) which follow one of three ARTs: heavy bourgeois males defend harems of females and have 10 times higher reproductive success than smaller roamers, which have ten times higher reproductive success than philopatric males, which remain in their natal group and are the smallest males. Bourgeois and philopatric males live in social groups that defend one territory, while roamers are solitary and roam over larger areas. We predicted that roamers will face higher energetic costs compared to group-living males because they do not gain thermoregulatory benefits of huddling in groups and might travel larger distances as they have larger home ranges. We measured daily energy expenditure (DEE) of 30 males, resting metabolic rate (RMR) of 79 males, travel distances and daily ranges of 31 males and changes in body mass of 51 males. Roamers had higher DEE and higher RMR than both types of group-living males. Philopatric males had shorter travel distances and smaller daily ranges than both roamers and bourgeois males, which did not differ from each other. This indicates that the higher DEE of roamers compared to bourgeois males cannot be explained by larger travel distances. Philopatrics gained body mass faster than bourgeois males and roamers, thereby increasing their competitive ability and thus the probability of later switching to a tactic of higher reproductive success. Our results suggest that roamers suffer energetic costs that might reduce their ability of gaining body mass and thus the likelihood of switching to the bourgeois tactic, indicating evolutionary trade-offs between investing energy into roaming versus gaining body mass.

Alternative male morphs solve sperm performance/longevity trade-off in opposite directions.

Males pursuing alternative reproductive tactics have been predicted to face a trade-off between maximizing either swimming performance or endurance of their sperm. However, empirical evidence for this trade-off is equivocal, which may be due to simplistic assumptions. In the shell-brooding cichlid fish Lamprologus callipterus, two Mendelian male morphs compete for fertilization by divergent means: Bourgeois nest males ejaculate sperm, on average, about six times farther from the unfertilized ova than do parasitic dwarf males. This asymmetry is opposite to the usual situation, in which bourgeois males typically benefit from superior fertilization opportunities, suggesting that nest males' sperm should persist longer than dwarf male sperm. The assumed trade-off between sperm swimming performance and longevity predicts that, in turn, sperm of dwarf males should outperform that of nest males in swimming efficiency. Measurement of sperm performance and endurance reveals that dwarf male spermatozoa swim straighter initially than those of nest males, but their motility declines earlier and their velocity slows down more abruptly. Nest male sperm survives longer, which relates to a larger sperm head plus midpiece, implying more mitochondria. Thus, the trade-off between sperm performance and endurance is optimized in opposite directions by alternative male morphs. We argue that the relative success of alternative sperm performance strategies can be influenced strongly by environmental factors such as the time window between gamete release and fertilization, and the position of gamete release. This is an important yet little understood aspect of gametic adaptations to sperm competition.

Female Mimicry by Sneaker Males Has a Transcriptomic Signature in Both the Brain and the Gonad in a Sex-Changing Fish.

Phenotypic plasticity represents an elegant adaptive response of individuals to a change in their environment. Bluehead wrasses (Thalassoma bifasciatum) exhibit astonishing sexual plasticity, including female-to-male sex change and discrete male morphs that differ strikingly in behavior, morphology, and gonadal investment. Using RNA-seq transcriptome profiling, we examined the genes and physiological pathways underlying flexible behavioral and gonadal differences among female, dominant (bourgeois) male, and female-mimic (sneaker) male blueheads. For the first time in any organism, we find that female mimicry by sneaker males has a transcriptional signature in both the brain and the gonad. Sneaker males shared striking similarity in neural gene expression with females, supporting the idea that males with alternative reproductive phenotypes have "female-like brains." Sneaker males also overexpressed neuroplasticity genes, suggesting that their opportunistic reproductive strategy requires a heightened capacity for neuroplasticity. Bourgeois males overexpressed genes associated with socio-sexual behaviors (e.g., isotocin), but also neuroprotective genes and biomarkers of oxidative stress and aging, indicating a hitherto unexplored cost to these males of attaining the reproductively privileged position at the top of the social hierarchy. Our novel comparison of testicular transcriptomes in a fish with male sexual polymorphism associates greater gonadal investment by sneaker males with overexpression of genes involved in cell proliferation and sperm quality control. We propose that morphological female-mimicry by sneaker male teleosts entails pervasive downregulation of androgenesis genes, consistent with low androgen production in males lacking well-developed secondary sexual characters.

The evolution of strategic male mating effort in an information transfer framework.

Sperm competition theory predicts that males should use cues indicating the risk and intensity of sperm competition to tailor their sperm investment accordingly. Rival males are an important source of social information regarding sperm competition risk. However, revealing such information may not be in the rival males' interest. Here, we use a theoretical approach based on informed and uninformed games to investigate when information transfer about sperm competition risk to competitors is beneficial for a male, and when it is not. The results show that signalling to potential future mates that a female has already mated is beneficial when the signalling male has a sperm competition disadvantage, whereas it is unfavourable when the signaller has an advantage. The reason for this counterintuitive result is that the rival males' optimal response is to reduce sperm investment when the signaller has a disadvantage and, conversely, to increase investment when the signaller has an advantage. Furthermore, we analysed scenarios where males use alternative reproductive tactics. In this situation, signalling the awareness of sperm competition risk rarely pays; instead, it is beneficial to maintain an information advantage. Thus, it may be beneficial for bourgeois males to accept cuckoldry instead of revealing their sperm competition awareness to reproductive parasites. These results provide new insight into the evolution of communication between rivals in the context of sperm competition.

Fragile Minds and Vulnerable Souls: The Matter of Obscenity in Nineteenth-Century Germany. By Sarah L. Leonard. Material Texts. Edited by Roger Chartier et al.Philadelphia: University of Pennsylvania Press, 2015. Pp. viii+258. $55.00 (cloth); $55.00 (e-book).

<i>Fragile Minds and Vulnerable Souls: The Matter of Obscenity in Nineteenth-Century Germany</i>. By Sarah L. Leonard. Material Texts. Edited by Roger Chartier et al.Philadelphia: University of Pennsylvania Press, 2015. Pp. viii+258. $55.00 (cloth); $55.00 (e-book).

Sperm-limited males save ejaculates for future matings when competing with superior rivals

Adjusting ejaculates to sperm competition can lead to sperm limitation. Particularly in polygynous species, males may face a trade-off between investing sperm in current or future mating opportunities. The optimal sperm allocation decision should depend on the relative intensity of sperm competition experienced in a mating sequence. Here we asked how males respond to this trade-off in polygynous fish with alternative male mating tactics, intense sperm competition and sperm limitation. Large bourgeois males of the shell-brooding cichlid Lamprologus callipterus build nests consisting of empty snail shells, in which females spawn and raise offspring. During spawning, nest males release ejaculates into the shell opening. Genetically distinct, parasitic dwarf males enter shells during spawning to fertilize the eggs from inside the shell. These dwarf males were previously shown to be superior sperm competitors to nest males. Here we showed that when spawning with several females simultaneously, nest males reduced the spawning duration for each clutch and the number of ejaculations per female tended to decrease, reflecting sperm limitation. Experimental exposure of nest males to sperm competition with dwarf males reduced the number and duration of ejaculations by roughly half. Hence, when exposed to competition with a superior rival, nest males did not increase their sperm expenditure as predicted by sperm competition risk models, but in fact saved sperm for future mating opportunities as predicted by sperm competition intensity theory. This seems to be adaptive because of the considerable sperm demands in this species, which is partly due to their high degree of polygyny.

Androgen responses to reproductive competition of males pursuing either fixed or plastic alternative reproductive tactics.

Alternative reproductive tactics (ARTs), which can be plastic or fixed for life, may be characterized by distinct hormonal profiles. The relative plasticity hypothesis predicts flexible androgen regulation for adult males pursuing plastic tactics, but a less flexible regulation for males using a fixed tactic throughout life. Furthermore, androgen profiles may respond to changes in the social environment, as predicted by the social reciprocity models of hormone/behaviour interactions. The cichlid fish Lamprologus callipterus provides a rare opportunity to study the roles of androgens for male ARTs within a single species, because fixed and plastic ARTs coexist. We experimentally exposed males to competitors pursuing either the same or different tactics to test predictions of the relative plasticity and the social reciprocity models. Androgen profiles of different male types partly comply with predictions derived from the relative plasticity hypothesis: males of the plastic bourgeois/sneaker male trajectory showed different 11-ketotestosterone (11-KT) levels when pursuing either bourgeois or parasitic sneaker male behaviours. Surprisingly, males pursuing the fixed dwarf male tactic showed the highest free and conjugated 11-KT and testosterone (T) levels. Our experimental social challenges significantly affected the free 11-KT levels of bourgeois males, but the androgen responses did not differ between challenges involving different types of competitors. Furthermore, the free T-responses of the bourgeois males correlated with their aggressive behaviour exhibited against competitors. Our results provide new insights into the endocrine responsiveness of fixed and plastic ARTs, confirming and refuting some predictions of both the relative plasticity and the social reciprocity models.

Contest versus scramble competition among males pursuing fixed or plastic alternative reproductive tactics

Reproductive and agonistic behaviours typically diverge between individuals pursuing alternative reproductive tactics (ARTs). When tactics are fixed for life, evolutionary theory predicts that the relative frequencies of alternative male genotypes are stabilized in a population by negative frequency dependence. This implies that competition is greatest between males pursuing the same tactic. The cichlid fish Lamprologus callipterus exhibits three male ARTs involving fixed and flexible tactics, and an extreme intrasexual size dimorphism determined by Mendelian inheritance. Large nest males defend territories and construct nests of empty snail shells in which females breed. In contrast, dwarf males pursuing a genetically fixed parasitic tactic enter shells surreptitiously during spawning in order to steal fertilizations. Sneaker males using another parasitic tactic, which is plastic and conditional, steal fertilizations opportunistically during spawning by quick intrusions into the nest. The variation in tactic origin and reproductive behaviour and the substantial asymmetry in body size render L. callipterus an ideal model system to study theoretical predictions regarding the types and intensity of contest behaviours among conspecific competitors pursuing ARTs. In an experiment exposing males to competitors using either the same or a different tactic, within-tactic competition was much more intense than between-tactic competition in bourgeois males, as predicted by evolutionary theory. In addition, the level of aggression displayed by bourgeois males against male intruders was apparently triggered by perceived differences in body size. In contrast to bourgeois males, parasitic males showed hardly any aggressive behaviour against other males, indicating that their contests follow the pattern of scramble competition. The conditions characterizing parasitic reproduction apparently select for rapid responsiveness when opportunities arise to fertilize eggs, whereas attacking other males in this situation seems inappropriate. Our results show that males pursuing ARTs diverge in the way they react to reproductive competition, mainly dependent on their overall resource defence strategy.

Alternative reproductive tactics in snail shell-brooding cichlids diverge in energy reserve allocation.

Life history theory predicts that the amount of resources allocated to reproduction should maximize an individual's lifetime reproductive success. So far, resource allocation in reproduction has been studied mainly in females. Intraspecific variation of endogenous energy storage and utilization patterns of males has received little attention, although these patterns may vary greatly between individuals pursuing alternative reproductive tactics (ARTs). ARTs are characterized by systematic variation of behavioral, physiological, and often morphological traits among same-sex conspecifics. Some individuals may rely on previously accumulated reserves, because of limited foraging opportunities during reproduction. Others may be able to continue foraging during reproduction, thus relying on reserves to a lesser extent. We therefore predicted that, if male tactics involve such divergent limitations and trade-offs within a species, ARTs should correspondingly differ in energy reserve allocation and utilization. To test this prediction, we studied short-term and long-term reserve storage patterns of males in the shell-brooding cichlid Lamprologus callipterus. In this species, bourgeois males investing in territory defense, courtship, and guarding of broods coexist with two distinct parasitic male tactics: (1) opportunistic sneaker males attempting to fertilize eggs by releasing sperm into the shell opening when a female is spawning; and (2) specialized dwarf males attempting to enter the shell past the spawning female to fertilize eggs from inside the shell. Sneaker males differed from other male types by showing the highest amount of accumulated short-term and long-term fat stores, apparently anticipating their upcoming adoption of the nest male status. In contrast, nest males depleted previously accumulated energy reserves with increasing nest holding period, as they invest heavily into costly reproductive behaviors while not taking up any food. This conforms to a capital breeder strategy. Dwarf males did not accumulate long-term fat stores at all, which they can afford due to their small behavioral effort during reproduction and their continued feeding activity, conforming to an income breeder strategy. Our data confirm that the resource storage patterns of males pursuing ARTs can diverge substantially, which adds to our understanding of the coexistence and maintenance of alternative reproductive patterns within species.

Y-linked Mendelian inheritance of giant and dwarf male morphs in shell-brooding cichlids

Behavioural variation among conspecifics is typically contingent on individual state or environmental conditions. Sex-specific genetic polymorphisms are enigmatic because they lack conditionality, and genes causing adaptive trait variation in one sex may reduce Darwinian fitness in the other. One way to avoid such genetic antagonism is to control sex-specific traits by inheritance via sex chromosomes. Here, controlled laboratory crossings suggest that in snail-brooding cichlid fish a single locus, two-allele polymorphism located on a sex-linked chromosome of heterogametic males generates an extreme reproductive dimorphism. Both natural and sexual selection are responsible for exceptionally large body size of bourgeois males, creating a niche for a miniature male phenotype to evolve. This extreme intrasexual dimorphism results from selection on opposite size thresholds caused by a single ecological factor, empty snail shells used as breeding substrate. Paternity analyses reveal that in the field parasitic dwarf males sire the majority of offspring in direct sperm competition with large nest owners exceeding their size more than 40 times. Apparently, use of empty snail shells as breeding substrate and single locus sex-linked inheritance of growth are the major ecological and genetic mechanisms responsible for the extreme intrasexual diversity observed in Lamprologus callipterus.

Sneaker males are not necessarily similar to females in colour in a sexually monochromatic cichlid

In species with alternative reproductive tactics (ARTs), males employing different tactics usually have different appearance. The clearest difference is body size: bourgeois males that monopolize access to females are larger than sneaker males that steal fertilizations from them. Sneakers are also known to be often dull in colour compared with bourgeois males and rather resemble females. However, this typical colour pattern is unlikely in the Lake Tanganyika cichlid Neolamprologus mondabu: we observed sneaking by two distinctive colour morphs, namely, black (which is apparently conspicuous against the background) and white (which is apparently background-matching). Because breeding females are black, this observation indicates that one type of sneakers contrasts female appearance. In this study, we conducted field studies to determine the expression of body colour in relation to ARTs in this fish. Body colour was dependent on size, but not sex and maturation, with black individuals being larger than white ones. Together with the presence of intermediate grey colour expressed by intermediate-sized individuals, this fish could ontogenetically changes their body colour from white to black. Both sexes of black individuals occupied feeding territories, but white individuals were non-territorial, indicating that the black body signals the possession of a feeding territory. Sexually active females were invariably black, whereas sexually active males were both black and white in colour. Few of the largest black males held harems, which included several female territories, whereas the remaining males were bachelors with no female territories. These bachelor males invested more in testes than harem males, suggesting that bachelors employ sneaking tactics, which is corroborated by our sneaking observations. To our knowledge, this is the first study showing that sneakers are entirely dissimilar to females in appearance. Herein, we discuss why sneakers are dichromatic in relation to their life histories.

Fellow men: Fantin-Latour and the problem of the group in nineteenth-century French painting

Near the end of her new study on the group portraits of Henri Fantin-Latour, Bridget Alsdorf notes: “The history of nineteenth-century French art is a field fascinated by movements and collective politics, yet still dominated by accounts of singular artists and oeuvres. Although we depend on groups to give structure to history, as artists depended on them to provide camaraderie and support, it has proved difficult to imagine the artistic self as formed fundamentally by way of others” (227). Her book Fellow Men: Fantin-Latour and the Problem of the Group in Nineteenth-Century French Painting goes a long way toward rectifying this scholarly imbalance. As her subtitle indicates, Alsdorf treats the group as a problem in more than one sense: she approaches it both from the perspective of representation, through a series of close readings of multi-figure portraits by Fantin and his contemporaries, and as a social, and indeed gender, issue relating to the dynamics of male interaction in the mid-nineteenth century. At the heart of her argument is a meditation on the necessary tensions generated by the conflicting pressures of mutuality and individualism—by the opposing needs for progressive artists to assert a group identity in the face of an indifferent or hostile public while at the same time cultivating the distinct creative personality increasingly demanded in the cultural marketplace. Alsdorf reveals traces of these tensions in the visual structures of the works she so persuasively explores, and gestures toward a broader analysis of their centrality in shaping individuals and groups within the artistic community, and among bourgeois males of the period more generally.

The same old songs in Reagan-era teen film

This article examines the recontextualization of 1950s rock in the form of “Oldies” in teen films of the 1970s and 1980s. Specifically, the article highlights the peculiar phenomenon of scenes featuring teenagers lip-synching to oldies songs in films like Risky Business (1983), Pretty in Pink (1986), Ferris Bueller's Day Off (1986), and Adventures in Babysitting (1987).In these scenes, like in the cover versions of rhythm and blues records popularized by white artists in the fifties, white teens embody black cultural forms, “covering” over the racial and sexual politics that characterized rock and roll's emergence. The transformation of rock 'n' roll from “race music” to the safe alternative for white bourgeois males in the face of new wave, punk, disco and hip hop, reflected in the establishment of oldies radio formats and revival tours, was aided and abetted by oldies soundtracks to Hollywood film.

Personality traits, reproductive behaviour and alternative mating tactics in male European bitterling, Rhodeus amarus

Individual differences in behavioural traits may play a role in reproductive behaviour and it is likely that different personality types have different reproductive success across specific social environments. This suggests a role for sexual selection for personality types, including a link between behavioural traits and alternative reproductive tactics. While research on morphological differences between guarders (bourgeois males defending resources) and sneakers (males parasitizing the territories of bourgeois males) is well characterized, the role of personality in the adoption of alternative mating tactics has hitherto been largely ignored. We investigated individual behavioural differences in a sneaker‐guarder mating system using size/age-matched males of a small freshwater fish, the European bitterling (Rhodeus amarus). We predicted distinct behavioural responses by guarder and sneaker males that were consistent in different contexts, indicating the existence of behavioural syndromes associated with male mating tactic. No behavioural syndromes were detected in male R. amarus, despite ability of individual male bitterling to establish dominance, boldness and investment in sperm competition being relatively repeatable across three consecutive trials. Male aggression, though not repeatable, was negatively correlated with the number of ejaculations, indicating a trade-off between aggression and sperm loading. No association between the tendency of males to guard a territory and behavioural traits was found, despite a significant association between the tendency to guard and morphological and physiological traits, with higher relative testis size and more breeding tubercles in guarder males. Our data suggest male bitterling mating tactics are largely unconstrained by innate factors and likely to be the product of prevailing environmental and social conditions.

Parker’s sneak-guard model revisited: why do reproductively parasitic males heavily invest in testes?

Alternative reproductive tactics are widespread in males and may cause intraspecific differences in testes investment. Parker's sneak-guard model predicts that sneaker males, who mate under sperm competition risk, invest in testes relatively more than bourgeois conspecifics that have lower risk. Given that sneakers are much smaller than bourgeois males, sneakers may increase testes investment to overcome their limited sperm productivity because of their small body sizes. In this study, we examined the mechanism that mediates differential testes investment across tactics in the Lake Tanganyika cichlid fish Lamprologus callipterus. In the Rumonge population of Burundi, bourgeois males are small compared with those in other populations and have a body size close to sneaky dwarf males. Therefore, if differences in relative testis investment depend on sperm competition, the rank order of relative testis investment should be dwarf males > bourgeois males in Rumonge = bourgeois males in the other populations. If differences in relative testis investment depend on body size, the rank order of relative testes investment should be dwarf males > bourgeois males in Rumonge > bourgeois males in the other populations. Comparisons of relative testis investment among the three male groups supported the role of sperm competition, as predicted by the sneak-guard model. Nevertheless, the effects of absolute body size on testes investment should be considered to understand the mechanisms underlying intraspecific variation in testes investment caused by alternative reproductive tactics.

Sperm phenotypic plasticity in a cichlid: a territorial male's counterstrategy to spawning takeover

Studies of sperm competition in species with alternative reproductive tactics (ARTs) often pay attention to the differences in investments in sperm between sneakers facing a higher sperm competition risk and bourgeois males facing a lower risk. Here, we examined within-tactic as well as among-tactic variations in sperm investments in the Lake Tanganyika shell-brooding cichlid Telmatochromis vittatus, a species with 2 types of parasitic tactics. Territorial male T. vittatus suffer reproductive parasitism by both smaller sneaker males and larger pirate males (‘‘pirates’’ take over the spawning event during which territorial males perform sneaking as a counterstrategy). We hypothesized that both territorial males living under the risk of pirates and sneakers face increased risk of sperm competition and therefore should produce high-quality sperm compared with both territorial males that experienced no piracy risk and pirates. As expected, field studies showed that the former 2 males produced longer lived sperm than the other males. Aquarium experiments demonstrated that a visual stimulus of a pirate was enough to induce an increase in sperm longevity in territorial males compared with when no such stimulus was given. These results indicate that territorial male T. vittatus can plastically adjust at least one sperm quality trait in response to piracy risk. Moreover, long-term monitoring of males in the field showed that small territorial males grow into large territorial males and finally into pirates, so ARTs are not fixed over life. Accordingly, male T. vittatus appear to ontogenetically change their sperm longevity in response to size-dependent sperm

Reproductive parasitism: male and female responses to conspecific and heterospecific intrusions at spawning in a mouth‐brooding cichlid Ophthalmotilapia ventralis

A rare form of alternative reproductive behaviour without simultaneous parasitic spawning was observed in Ophthalmotilapia ventralis, a lekking mouth-brooding cichlid from Lake Tanganyika. Floater males attempted to sneak opportunistically into the territory to actively court the female, while the owner (bourgeois male) defended the territory against other potential intruders. Floater males had more body fat than territory owners and generally higher condition factors. In field experiments, the response of bourgeois males and courted females was tested towards floaters and egg predators (a catfish Synodontis multipunctatus) present in the territories. Territory owners responded aggressively particularly to floaters, and female responsiveness to bourgeois male courtship tended to decline when floaters were present. The potential influence of reproductive parasitism on sexual selection in mouth-brooding cichlids is discussed.

COMPARATIVE PHYLOGENETIC ANALYSIS OF MALE ALTERNATIVE REPRODUCTIVE TACTICS IN RAY-FINNED FISHES

Using comparative phylogenetic analysis, we analyzed the evolution of male alternative reproductive tactics (MARTs) in ray-finned fishes (Actinopterygii). Numerous independent origins for each type of MART (involving sneaker males, female mimics, pirates, and satellite males) indicate that these behaviors have been highly labile across actinopterygiian evolution, consistent with a previous notion that convergent selection in fishes can readily mold the underlying suites of reproductive hormones into similar behaviors. The evolutionary appearance of MARTs was significantly correlated with the presence of sexually selected traits in bourgeois males (P = 0.001) but not with the presence of male parental care. This suggests that MARTs often arise from selection on some males to circumvent bourgeois male investment in mate monopolization, rather than to avoid male brood care per se. We found parsimony evidence for an evolutionary progression of MARTs wherein sneaking is usually the evolutionary precursor to the presumably more complex MARTs of female mimicry and cooperative satellite behavior. Nest piracy appears not to be part of this evolutionary progression, possibly because its late onset in the life cycle of most ray-finned fishes reduces the effects of selection on this reproductive tactic.