Scorpionfish rapidly change colour in response to their background.

BACKGROUND: To facilitate background matching in heterogenous environments, some animals rapidly change body colouration. Marine predatory fishes might use this ability to hide from predators and prey. Here, we focus on scorpionfishes (Scorpaenidae), well-camouflaged, bottom-dwelling sit-and-wait predators. We tested whether Scorpaena maderensis and Scorpaena porcus adjust body luminance and hue in response to three artificial backgrounds and thereby achieve background matching. Both scorpionfish species are also red fluorescent, which could contribute to background matching at depth. Therefore, we tested whether red fluorescence is also regulated in response to different backgrounds. The darkest and the lightest backgrounds were grey, while the third background was orange of intermediate luminance. Scorpionfish were placed on all three backgrounds in a randomised repeated measures design. We documented changes in scorpionfish luminance and hue with image analysis and calculated contrast to the backgrounds. Changes were quantified from the visual perspective of two potential prey fishes, the triplefin Tripterygion delaisi and the goby Pomatoschistus flavescens. Additionally, we measured changes in the area of scorpionfish red fluorescence. Because scorpionfish changed quicker than initially expected, we measured luminance change at a higher temporal resolution in a second experiment. RESULTS: Both scorpionfish species rapidly adjusted luminance and hue in response to a change of background. From prey visual perspective, scorpionfishes' body achromatic and chromatic contrasts against the background were high, indicating imperfect background matching. Chromatic contrasts differed considerably between the two observer species, highlighting the importance of choosing natural observers with care when studying camouflage. Scorpionfish displayed larger areas of red fluorescence with increasing luminance of the background. With the second experiment, we showed that about 50% of the total luminance change observed after one minute is achieved very rapidly, in five to ten seconds. CONCLUSION: Both scorpionfish species change body luminance and hue in response to different backgrounds within seconds. While the achieved background matching was suboptimal for the artificial backgrounds, we propose that the observed changes were intended to reduce detectability, and are an essential strategy to camouflage in the natural environment.

The round goby genome provides insights into mechanisms that may facilitate biological invasions.

BACKGROUND: The invasive benthic round goby (Neogobius melanostomus) is the most successful temperate invasive fish and has spread in aquatic ecosystems on both sides of the Atlantic. Invasive species constitute powerful in situ experimental systems to study fast adaptation and directional selection on short ecological timescales and present promising case studies to understand factors involved the impressive ability of some species to colonize novel environments. We seize the unique opportunity presented by the round goby invasion to study genomic substrates potentially involved in colonization success. RESULTS: We report a highly contiguous long-read-based genome and analyze gene families that we hypothesize to relate to the ability of these fish to deal with novel environments. The analyses provide novel insights from the large evolutionary scale to the small species-specific scale. We describe expansions in specific cytochrome P450 enzymes, a remarkably diverse innate immune system, an ancient duplication in red light vision accompanied by red skin fluorescence, evolutionary patterns of epigenetic regulators, and the presence of osmoregulatory genes that may have contributed to the round goby's capacity to invade cold and salty waters. A recurring theme across all analyzed gene families is gene expansions. CONCLUSIONS: The expanded innate immune system of round goby may potentially contribute to its ability to colonize novel areas. Since other gene families also feature copy number expansions in the round goby, and since other Gobiidae also feature fascinating environmental adaptations and are excellent colonizers, further long-read genome approaches across the goby family may reveal whether gene copy number expansions are more generally related to the ability to conquer new habitats in Gobiidae or in fish.

Redirection of ambient light improves predator detection in a diurnal fish.

Cases where animals use controlled illumination to improve vision are rare and thus far limited to chemiluminescence, which only functions in darkness. This constraint was recently relaxed by studies on Tripterygion delaisi, a small triplefin that redirects sunlight instead. By reflecting light sideways with its iris, it has been suggested to induce and detect eyeshine in nearby micro-prey. Here, we test whether 'diurnal active photolocation' also improves T. delaisi's ability to detect the cryptobenthic sit-and-wait predator Scorpaena porcus, a scorpionfish with strong daytime retroreflective eyeshine. Three independent experiments revealed that triplefins in which light redirection was artificially suppressed approached scorpionfish significantly closer than two control treatments before moving away to a safer distance. Visual modelling confirmed that ocular light redirection by a triplefin is sufficiently strong to generate a luminance increase in scorpionfish eyeshine that can be perceived by the triplefin over 6-8 cm under average conditions. These distances coincide well with the closest approaches observed. We conclude that light redirection by small, diurnal fish significantly contributes to their ability to visually detect cryptic predators, strongly widening the conditions under which active sensing with light is feasible. We discuss the consequences for fish eye evolution.

Small benthic fish strike at prey over distances that fall within theoretical predictions for active sensing using light.

Small, benthic, cryptic fishes represent a species-rich guild on marine substrates. Most of them are micropredators that feed on crustaceans that are often smaller than 1 mm. Typical examples are seahorses and pipefishes (Syngnathidae), most gobies (Gobiidae), dragonets (Callionymidae) and triplefins (Tripterygiidae). Previous work on the yellow black-headed triplefin Tripterygion delaisi demonstrated that it actively redirects downwelling sunlight sideways using its iris and can use this to locally illuminate objects of interest. We call this form of active sensing using light "diurnal active photolocation". Visual modelling predicted that light redirection can be sufficient to induce a perceivable change in luminance in the eyes of one of its prey species, a cryptic gammarid crustacean (Cheirocratus sp.), over distances of 1-2 cm. Empirical validation, however, was not possible because measurements of predation distances have not been quantified for free-ranging, small, benthic fishes before. Here, we present interaction distances measured from videos of T. delaisi approaching and striking at prey in the field. Out of 160 recordings, we were able to quantify 78 prey approaching distances and 100 striking distances. Approaching distances ranged from 2.1 to 4.1 cm (interquartile range, IQR) and involved one to five approaching steps before the actual strike occurred. The distance over which the final strike took place varied from 0.7 to 1.6 cm (IQR). Both approaching and striking distances increased with fish body size. We conclude that most approaching sequences started too far away to be explained by prey detection through light redirection. Striking distances, however, fell well with the distances predicted by the model. We conclude that if diurnal active photolocation plays a role in prey detection, it is during the final decision whether to strike or not.

The contrast sensitivity function of a small cryptobenthic marine fish.

Spatial resolution is a key property of eyes when it comes to understanding how animals' visual signals are perceived. This property can be robustly estimated by measuring the contrast sensitivity as a function of different spatial frequencies, defined as the number of achromatic vertical bright and dark stripe pairs within one degree of visual angle. This contrast sensitivity function (CSF) has been estimated for different animal groups, but data on fish are limited to two free-swimming, freshwater species (i.e., goldfish and bluegill sunfish). In this study, we describe the CSF of a small marine cryptobenthic fish (Tripterygion delaisi) using an optokinetic reflex approach. Tripterygion delaisi features a contrast sensitivity that is as excellent as other fish species, up to 125 (reciprocal of Michelson contrast) at the optimal spatial frequency of 0.375 c/°. The maximum spatial resolution is instead relatively coarse, around 2.125 c/°. By comparing our results with acuity values derived from anatomical estimates of ganglion cells' density, we conclude that the optokinetic reflex seems to be adapted to process low spatial frequency information from stimuli in the peripheral visual field and show that small marine fish can feature excellent contrast sensitivity at optimal spatial frequency.

Optic-nerve-transmitted eyeshine, a new type of light emission from fish eyes.

BACKGROUND: Most animal eyes feature an opaque pigmented eyecup to assure that light can enter from one direction only. We challenge this dogma by describing a previously unknown form of eyeshine resulting from light that enters the eye through the top of the head and optic nerve, eventually emanating through the pupil as a narrow beam: the Optic-Nerve-Transmitted (ONT) eyeshine. We characterize ONT eyeshine in the triplefin blenny Tripterygion delaisi (Tripterygiidae) in comparison to three other teleost species, using behavioural and anatomical observations, spectrophotometry, histology, and magnetic resonance imaging. The study's aim is to identify the factors that determine ONT eyeshine occurrence and intensity, and whether these are specifically adapted for that purpose. RESULTS: ONT eyeshine intensity benefits from locally reduced head pigmentation, a thin skull, the gap between eyes and forebrain, the potential light-guiding properties of the optic nerve, and, most importantly, a short distance between the head surface and the optic nerves. CONCLUSIONS: The generality of these factors and the lack of specifically adapted features implies that ONT eyeshine is widespread among small fish species. Nevertheless, its intensity varies considerably, depending on the specific combination and varying expression of common anatomical features. We discuss whether ONT eyeshine might affect visual performance, and speculate about possible functions such as predator detection, camouflage, and intraspecific communication.

Fluorescence characterisation and visual ecology of pseudocheilinid wrasses.

BACKGROUND: Wrasses represent the second largest family of marine fishes and display a high diversity of complex colours linked to ecological functions. Recently, red autofluorescent body colouration has been reported in some of these fishes. However, little is known about the distribution of such fluorescent body patterns in wrasses or the animals' ability to perceive such colours. RESULTS: Against this background, we (1) investigated long-wavelength emission autofluorescence in thirteen species of pseudocheilinid wrasses and (2) characterised the spectral absorbance of visual pigments in one of the examined species, the fairy wrasse Cirrhilabrus solorensis. Spectrophotometric analysis revealed that fluorescent body colouration is widespread and diverse within this clade, with considerable variation in both fluorescent pattern and maximum emission wavelength between species. Characterisation of visual pigments in retinal photoreceptors showed a single class of rod and three spectrally distinct cone photoreceptors, suggesting possible trichromacy. CONCLUSION: Combining the emission characteristics of fluorescence body colouration and the spectral sensitivity data of retinal cells suggests that the visual system of C. solorensis is sensitive to pseudocheilinid fluorescence.

Fairy wrasses perceive and respond to their deep red fluorescent coloration.

Fluorescence enables the display of wavelengths that are absent in the natural environment, offering the potential to generate conspicuous colour contrasts. The marine fairy wrasse Cirrhilabrus solorensis displays prominent fluorescence in the deep red range (650-700 nm). This is remarkable because marine fishes are generally assumed to have poor sensitivity in this part of the visual spectrum. Here, we investigated whether C. solorensis males can perceive the fluorescence featured in this species by testing whether the presence or absence of red fluorescence affects male-male interactions under exclusive blue illumination. Given that males respond aggressively towards mirror-image stimuli, we quantified agonistic behaviour against mirrors covered with filters that did or did not absorb long (i.e. red) wavelengths. Males showed significantly fewer agonistic responses when their fluorescent signal was masked, independent of brightness differences. Our results unequivocally show that C. solorensis can see its deep red fluorescent coloration and that this pattern affects male-male interactions. This is the first study to demonstrate that deep red fluorescent body coloration can be perceived and has behavioural significance in a reef fish.

Red fluorescence increases with depth in reef fishes, supporting a visual function, not UV protection.

Why do some marine fishes exhibit striking patterns of natural red fluorescence? In this study, we contrast two non-exclusive hypotheses: (i) that UV absorption by fluorescent pigments offers significant photoprotection in shallow water, where UV irradiance is strongest; and (ii) that red fluorescence enhances visual contrast at depths below -10 m, where most light in the 'red' 600-700 nm range has been absorbed. Whereas the photoprotection hypothesis predicts fluorescence to be stronger near the surface and weaker in deeper water, the visual contrast hypothesis predicts the opposite. We used fluorometry to measure red fluorescence brightness in vivo in individuals belonging to eight common small reef fish species with conspicuously red fluorescent eyes. Fluorescence was significantly brighter in specimens from the -20 m sites than in those from -5 m sites in six out of eight species. No difference was found in the remaining two. Our results support the visual contrast hypothesis. We discuss the possible roles fluorescence may play in fish visual ecology and highlight the possibility that fluorescent light emission from the eyes in particular may be used to detect cryptic prey.

Regulation of red fluorescent light emission in a cryptic marine fish.

INTRODUCTION: Animal colouration is a trade-off between being seen by intended, intra- or inter-specific receivers while not being seen by the unintended. Many fishes solve this problem by adaptive colouration. Here, we investigate whether this also holds for fluorescent pigments. In those aquatic environments in which the ambient light is dominated by bluish light, red fluorescence can generate high-contrast signals. The marine, cryptic fish Tripterygion delaisi inhabits such environments and has a bright red-fluorescent iris that can be rapidly up- and down-regulated. Here, we described the physiological and cellular mechanism of this phenomenon using a neurostimulation treatment with KCl and histology. RESULTS: KCl-treatment revealed that eye fluorescence regulation is achieved through dispersal and aggregation of black-pigmented melanosomes within melanophores. Histology showed that globular, fluorescent iridophores on the anterior side of the iris are grouped and each group is encased by finger-like extensions of a single posterior melanophore. Together they form a so-called chromatophore unit. By dispersal and aggregation of melanosomes into and out of the peripheral membranous extensions of the melanophore, the fluorescent iridophores are covered or revealed on the anterior (outside) of the iris. CONCLUSION: T. delaisi possesses a well-developed mechanism to control the fluorescent emission from its eyes, which may be advantageous given its cryptic lifestyle. This is the first time chromatophore units are found to control fluorescent emission in marine teleost fishes. We expect other fluorescent fish species to use similar mechanisms in the iris or elsewhere in the body. In contrast to a previously described mechanism based on dendritic fluorescent chromatophores, chromatophore units control fluorescent emission through the cooperation between two chromatophore types: an emitting and an occluding type. The discovery of a second mechanism for fluorescence modulation strengthens our view that fluorescence is a relevant and adaptive component of fish colouration.

Scorpionfish adjust skin pattern contrast on different backgrounds.

The two scorpionfish species Scorpaena maderensis and S. porcus are well camouflaged ambush predators that rapidly change body colouration to adjust to background colour in less than 1 min. We tested whether individuals of both species also adjust body pattern to that of the background. We placed fish on backgrounds of different pattern granularity and quantified the change in fish body pattern over 1 min. We used calibrated image analysis to analyse the patterns from the visual perspective of a prey fish species using a granularity (pattern energy) analysis and an image clustering approach. In our experiment, fish did not change their most contrasting pattern components as defined by the dominant marking size, but changed their average marking size. Moreover, fish responded with a change in pattern in contrast to the different experimental backgrounds, especially when compared to the acclimation phase. These results indicate that scorpionfish have one main pattern that can be adjusted by modulating its internal contrast. A reduction in pattern contrast could thereby improve background matching, while an increase could promote camouflage via disruptive colouration.

Sex differences in host defence interfere with parasite-mediated selection for outcrossing during host-parasite coevolution.

The Red Queen hypothesis proposes that coevolving parasites select for outcrossing in the host. Outcrossing relies on males, which often show lower immune investment due to, for example, sexual selection. Here, we demonstrate that such sex differences in immunity interfere with parasite-mediated selection for outcrossing. Two independent coevolution experiments with Caenorhabditis elegans and its microparasite Bacillus thuringiensis produced decreased yet stable frequencies of outcrossing male hosts. A subsequent systematic analysis verified that male C. elegans suffered from a direct selective disadvantage under parasite pressure (i.e. lower resistance, decreased sexual activity, increased escape behaviour), which can reduce outcrossing and thus male frequencies. At the same time, males offered an indirect selective benefit, because male-mediated outcrossing increased offspring resistance, thus favouring male persistence in the evolving populations. As sex differences in immunity are widespread, such interference of opposing selective constraints is likely of central importance during host adaptation to a coevolving parasite.

Multiple reciprocal adaptations and rapid genetic change upon experimental coevolution of an animal host and its microbial parasite.

The coevolution between hosts and parasites is predicted to have complex evolutionary consequences for both antagonists, often within short time periods. To date, conclusive experimental support for the predictions is available mainly for microbial host systems, but for only a few multicellular host taxa. We here introduce a model system of experimental coevolution that consists of the multicellular nematode host Caenorhabditis elegans and the microbial parasite Bacillus thuringiensis. We demonstrate that 48 host generations of experimental coevolution under controlled laboratory conditions led to multiple changes in both parasite and host. These changes included increases in the traits of direct relevance to the interaction such as parasite virulence (i.e., host killing rate) and host resistance (i.e., the ability to survive pathogens). Importantly, our results provide evidence of reciprocal effects for several other central predictions of the coevolutionary dynamics, including (i) possible adaptation costs (i.e., reductions in traits related to the reproductive rate, measured in the absence of the antagonist), (ii) rapid genetic changes, and (iii) an overall increase in genetic diversity across time. Possible underlying mechanisms for the genetic effects were found to include increased rates of genetic exchange in the parasite and elevated mutation rates in the host. Taken together, our data provide comprehensive experimental evidence of the consequences of host-parasite coevolution, and thus emphasize the pace and complexity of reciprocal adaptations associated with these antagonistic interactions.

Increased responsiveness in feeding behaviour of Caenorhabditis elegans after experimental coevolution with its microparasite Bacillus thuringiensis.

Immune responses, either constitutive or induced, are costly. An alternative defence strategy may be based on behavioural responses. For example, avoidance behaviour reduces contact with pathogens and thus the risk of infection as well as the requirement of immune system activation. Similarly, if pathogens are taken up orally, preferential feeding of pathogen-free food may be advantageous. Behavioural defences have been found in many animals, including the nematode Caenorhabditis elegans. We here tested nematodes from a laboratory based evolution experiment which had either coevolved with their microparasite Bacillus thuringiensis (BT) or evolved under control conditions. After 48 generations, coevolved populations were more sensitive to food conditions: in comparison with the controls, they reduced feeding activity in the presence of pathogenic BT strains while at the same time increasing it in the presence of non-pathogenic strains. We conclude that host-parasite coevolution can drive changes in the behavioural responsiveness to bacterial microbes, potentially leading to an increased defence against pathogens.

Host-parasite local adaptation after experimental coevolution of Caenorhabditis elegans and its microparasite Bacillus thuringiensis.

Coevolving hosts and parasites can adapt to their local antagonist. In studies on natural populations, the observation of local adaptation patterns is thus often taken as indirect evidence for coevolution. Based on this approach, coevolution was previously inferred from an overall pattern of either parasite or host local adaptation. Many studies, however, failed to detect such a pattern. One explanation is that the studied system was not subject to coevolution. Alternatively, coevolution occurred, but remained undetected because it took different routes in different populations. In some populations, it is the host that is locally adapted, whereas in others it is the parasite, leading to the absence of an overall local adaptation pattern. Here, we test for overall as well as population-specific patterns of local adaptation using experimentally coevolved populations of the nematode Caenorhabditis elegans and its bacterial microparasite Bacillus thuringiensis. Furthermore, we assessed the importance of random interaction effects using control populations that evolved in the absence of the respective antagonist. Our results demonstrate that experimental coevolution produces distinct local adaptation patterns in different replicate populations, including host, parasite or absence of local adaptation. Our study thus provides experimental evidence of the predictions of the geographical mosaic theory of coevolution, i.e. that the interaction between parasite and host varies across populations.

Control of social monogamy through aggression in a hermaphroditic shrimp.

INTRODUCTION: Sex allocation theory predicts that in small mating groups simultaneous hermaphroditism is the optimal form of gender expression. Under these conditions, male allocation is predicted to be very low and overall per-capita reproductive output maximal. This is particularly true for individuals that live in pairs, but monogamy is highly susceptible to cheating by both partners. However, certain conditions favour social monogamy in hermaphrodites. This study addresses the influence of group size on group stability and moulting cycles in singles, pairs, triplets and quartets of the socially monogamous shrimp Lysmata amboinensis, a protandric simultaneous hermaphrodite. RESULTS: The effect of group size was very strong: Exactly one individual in each triplet and exactly two individuals in each quartet were killed in aggressive interactions, resulting in group sizes of two individuals. All killed individuals had just moulted. No mortality occurred in single and pair treatments. The number of moults in the surviving shrimp increased significantly after changing from triplets and quartets to pairs. CONCLUSION: Social monogamy in L. amboinensis is reinforced by aggressive expulsion of supernumerous individuals. We suggest that the high risk of mortality in triplets and quartets results in suppression of moulting in groups larger than two individuals and that the feeding ecology of L. amboinensis favours social monogamy.

A context analysis of bobbing and fin-flicking in a small marine benthic fish.

Most antipredator strategies increase survival of individuals by signaling to predators, by reducing the chances of being recognized as prey, or by bewildering a predator's perception. In fish, bobbing and fin-flicking are commonly considered as pursuit-deterrent behaviors that signal a predator that it has been detected and thus lost its surprise-attack advantage. Yet, very few studies assessed whether such behavioral traits are restricted to the visual presence of a predator. In this study, we used the yellow black-headed triplefin Tripterygion delaisi to investigate the association between these behaviors and the visual exposure to (a) a black scorpionfish predator (Scorpaena porcus), (b) a stone of a size similar to that of S. porcus, (c) a conspecific, and (d) a harmless heterospecific combtooth blenny (Parablennius sanguinolentus). We used a laboratory-controlled experiment with freshly caught fish designed to test for differences in visual cues only. Distance kept by the focal fish to each stimulus and frequency of bobbing and fin-flicking were recorded. Triplefins kept greater distance from the stimulus compartment when a scorpionfish predator was visible. Bobbing was more frequent in the visual presence of a scorpionfish, but also shown toward the other stimuli. However, fin flicks were equally abundant across all stimuli. Both behaviors decreased in frequency over time suggesting that triplefin become gradually comfortable in a nonchanging new environment. We discuss why bobbing and fin-flicking are not exclusive pursuit-deterrent behaviors in this species, and propose additional nonexclusive functions such as enhancing depth perception by parallax motion (bobbing) or signaling vigilance (fin-flicking).

The function of copulatory plugs in Caenorhabditis remanei: hints for female benefits.

BACKGROUND: Mating plugs that males place onto the female genital tract are generally assumed to prevent remating with other males. Mating plugs are usually explained as a consequence of male-male competition in multiply mating species. Here, we investigated whether mating plugs also have collateral effects on female fitness. These effects are negative when plugging reduces female mating rate below an optimum. However, plugging may also be positive when plugging prevents excessive forced mating and keeps mating rate closer to a females' optimum. Here, we studied these consequences in the gonochoristic nematode Caenorhabditis remanei. We employed a new CO2-sedation technique to interrupt matings before or after the production of a plug. We then measured mating rate, attractiveness and offspring number. RESULTS: The presence of a mating plug did not affect mating rate or attractiveness to roving males. Instead, females with mating plugs produced more offspring than females without copulatory plugs. CONCLUSIONS: Our experiment suggests that plugging might have evolved under male-male competition but represents a poor protection against competing males in our experiment. Even if plugging does not reduce mating rate, our results indicate that females may benefit from being plugged in a different sense than remating prevention.

The costs and benefits of occasional sex: theoretical predictions and a case study.

Theory predicts that occasional sexual reproduction in predominantly parthenogenetic organisms offers all the advantages of obligate sexuality without paying its full costs. However, empirical examples identifying and evaluating the costs and benefits of rare sex are scarce. After reviewing the theoretical perspective on rare sex, we present our findings of potential costs and benefits of occasional sex in polyploid, sperm-dependent parthenogens of the planarian flatworm Schmidtea polychroa. Despite costs associated with the production of less fertile tetraploids as sexual intermediates, the benefits of rare sex prevail in S. polychroa and may be sufficiently strong to prevent extinction of parthenogenetic populations. This offers an explanation for the dominance of parthenogenesis in S. polychroa. We discuss the enigmatic question why not all organisms show a mixed reproduction mode.

No effect of mate novelty on sexual motivation in the freshwater snail Biomphalaria glabrata.

BACKGROUND: When mating effort (e.g. via ejaculates) is high, males are expected to strategically allocate their resources depending on the expected fitness gains from a given mating opportunity. One mechanism to achieve strategic mating is the Coolidge effect, where male sexual motivation declines across repeated encounters with a familiar partner, but resuscitates when encountering a novel female. Experimental tests of male mate choice via mechanisms such as the Coolidge effect, however, remain scarce. Moreover, it is untested to date whether the Coolidge effect occurs in a sex-specific manner in simultaneous hermaphrodites, where the motivation to mate with a familiar partner may vary with previous mating activity in the male or female role. RESULTS: We exposed focal hermaphroditic freshwater snails, Biomphalaria glabrata, repeatedly to either a familiar or a novel partner. None of our proxies of sexual motivation (remating likelihood, mating delay, copulation duration) varied between the novel and familiar partner treatments. Moreover, the mating role taken during the first copulation did not affect the subsequent choice of mating roles in the familiar partner treatment as would be expected if focals preferred to avoid mating twice in the same role with a familiar partner. This indicates the absence of sex-specific effects of partner novelty. CONCLUSION: Our data indicate that mate novelty does affect neither overall sexual motivation nor the choice of mating roles in B. glabrata. Hence, male mate choice via a Coolidge effect appears inexistent in this invertebrate hermaphrodite. We discuss the possible roles of insufficient fitness gains for discriminatory behaviour in populations with frequent mate encounters as well as poor mate discrimination capacities. Our findings lend also no support to the novel prediction that sexual motivation in simultaneous hermaphrodites varies with the mating roles taken during previous copulations, calling for empirical investigation in further hermaphrodite systems.