Mate Encounter Rates Research Articles

First evidence of sexual dimorphism in olfactory organs of deep-sea lanternfishes (Myctophidae).

Finding a mate is of the utmost importance for organisms, and the traits associated with successfully finding one can be under strong selective pressures. In habitats where biomass and population density is often low, like the enormous open spaces of the deep sea, animals have evolved many adaptations for finding mates. One convergent adaptation seen in many deep-sea fishes is sexual dimorphism in olfactory organs, where, relative to body size, males have evolved greatly enlarged olfactory organs compared to females. Females are known to give off chemical cues such as pheromones, and these chemical stimuli can traverse long distances in the stable, stratified water of the deep sea and be picked up by the olfactory organs of males. This adaptation is believed to help males in multiple lineages of fishes find mates in deep-sea habitats. In this study, we describe the first morphological evidence of sexual dimorphism in the olfactory organs of lanternfishes (Myctophidae) in the genus Loweina. Lanternfishes are one of the most abundant vertebrates in the deep sea and are hypothesized to use visual signals from bioluminescence for mate recognition or mate detection. Bioluminescent cues that are readily visible at distances as far as 10 m in the aphotic deep sea are likely important for high population density lanternfish species that have high mate encounter rates. In contrast, myctophids found in lower density environments where species encounter rates are lower, like those in Loweina, likely benefit from longer-range chemical or olfactory cues for finding and identifying mates.

Reproductive phenology and female mating frequency of the praying mantidTenodera angustipennisin western Japan

AbstractThe timing and frequency of female mating are important determinants of male reproductive success. Elucidating reproductive phenology is crucial to understand the evolution of mating behaviour and mating systems.Mate encounter rate is a key variable for understanding the evolutionary consequences of sexual cannibalism. However, remarkably little is known about female mating frequency in wild populations in mantids, charismatic insects that exhibit sexual cannibalism.The authors examined the reproductive phenology of a wild population of the sexually cannibalistic praying mantidTenodera angustipennis, and paid special attention to female mating frequency.Field surveys throughout two reproductive seasons were combined with survival model analysis to estimate the phenology of eclosion, adult sex ratio, female first mating, and oviposition, allowing quantification of time windows for reproductive maturation and female mating.Genetic paternity analysis using newly developed microsatellite markers revealed that females mated with two to six males on average before oviposition in the wild.The results provide a comprehensive record of the reproductive phenology and female mating frequency in a wild mantid population, and insight into the evolution of male mating behaviour under sperm competition and sexual cannibalism.

Black widows as plastic wallflowers: female choosiness increases with indicators of high mate availability in a natural population

Female choice is an important driver of sexual selection, but can be costly, particularly when choosy females risk remaining unmated or experience delays to reproduction. Thus, females should reduce choosiness when mate encounter rates are low. We asked whether choosiness is affected by social context, which may provide reliable information about the local availability of mates. This has been demonstrated in the lab, but rarely under natural conditions. We studied western black widow spiders (Latrodectus hesperus) in the field, placing experimental final-instar immature females so they were either ‘isolated’ or ‘clustered’ near naturally occurring conspecifics (≥10 m or ≤1 m, respectively, from a microhabitat occupied by at least one other female). Upon maturity, females in both treatments were visited by similar numbers of males, but clustered females were visited by males earlier and in more rapid succession than isolated females, confirming that proximity to conspecifics reduces the risk of remaining unmated. As predicted, isolated females were less choosy in staged mating trials, neither rejecting males nor engaging in pre-copulatory cannibalism, in contrast to clustered females. These results demonstrate that exposure of females to natural variation in demography in the field can alter choosiness of adults. Thus, female behaviour in response to cues of local population density can affect the intensity of sexual selection on males in the wild.

Mate-finding Allee effects can be exacerbated or relieved by sexual cannibalism.

Allee effects occur when individual or population survival decreases due to populations being small or sparse. A key mechanism underlying Allee effects is difficulty in finding mates at low densities. Species may be particularly vulnerable to mate-finding Allee effects if females rely on an abundance of males to reproduce successfully. In sexually cannibalistic species, females may consume males before or after copulation, potentially reducing the supply of males to the point where a mate-finding Allee effect occurs. In this study, we investigate the extent to which sexual cannibalism can modulate mate-finding Allee effects, and the conditions under which sexual cannibalism is likely to be particularly detrimental to population viability. We created an individual-based model that tracked specific females throughout the breeding season and used extinction risk and per capita growth rate to measure the strength of the Allee effects. We varied both founder population size and mate encounter rate independently of each other to expose the mechanism driving the Allee effects. We also analysed how cannibalism-derived female fecundity benefits affected extinction risk. We found that sexual cannibalism could lead to high extinction risk, particularly when cannibalism occurred before copulation, founder population size was small and mate encounter rates were low. However, post-copulatory cannibalism reduced extinction risk, if cannibalism increased female fecundity enough. We found that there were strong threshold effects, in which small changes in encounter rate could strongly alter population extinction risk. We find that sexual cannibalism is likely to negatively impact population survival as population size and mate encounter rate decrease. This may be exacerbated if male quality declines and female hunger increases in declining populations. As many top invertebrate predators, such as spiders and mantises, are sexually cannibalistic, this may have ecosystem-wide impacts. We also suggest that other reproductive behaviours, such as rejecting all but high-quality mates or requiring multiple mates to ensure fertility, are also likely to cause mate-finding Allee effects when habitat quality degrades.

The Evolution of Egg Trading in Simultaneous Hermaphrodites.

Egg trading-whereby simultaneous hermaphrodites exchange each other's eggs for fertilization-constitutes one of the few rigorously documented and most widely cited examples of direct reciprocity among unrelated individuals. Yet how egg trading may initially invade a population of nontrading simultaneous hermaphrodites is still unresolved. Here, we address this question with an analytical model that considers mate encounter rates and costs of egg production in a population that may include traders (who provide eggs for fertilization only if their partners also have eggs to reciprocate), providers (who provide eggs regardless of whether their partners have eggs to reciprocate), and withholders (cheaters who mate only in the male role and just use their eggs to elicit egg release from traders). Our results indicate that a combination of intermediate mate encounter rates, sufficiently high costs of egg production, and a sufficiently high probability that traders detect withholders (in which case eggs are not provided) is conducive to the evolution of egg trading. Under these conditions, traders can invade-and resist invasion from-providers and withholders alike. The prediction that egg trading evolves only under these specific conditions is consistent with the rare occurrence of this mating system among simultaneous hermaphrodites.

Population Structure, Movement Patterns, and Frequency of Multiple Matings in Tenodera sinensis (Mantodea: Mantidae).

Models of the evolution of sexual cannibalism show that the frequency of male mating opportunities has significant impact on male choice and male risk aversion. In this study, we examined ecological components that should affect opportunities for multiple mating in wild populations of the Chinese mantid (Tenodera sinensis Saussure). While conducting mark-recapture studies of two field populations over the course of two seasons, along with Global Positioning System data on locations of individuals, we collected data on population densities, movement patterns, and individual ranges to estimate the overlap of adult males and female mantids. Our results show that local populations of mantids range from 89 to 161 individuals and occur at densities ranging from 10 to 39 mantids per 1,000 m2. Males move greater distances daily compared with females, giving males larger home range sizes. The ranges of male mantids overlapped with multiple females, thus offering the potential for multiple mating by males. We directly observed 11 encounters between male and female T. sinensis, including one multiple mating by an individual male. The overall mate encounter rate for males was 12.5%. We also provide additional observations of interspecific sexual attraction between T. sinensis and Mantis religiosa Linne (Mantodea: Mantidae). Mantids were most commonly found within the top 20% of two flowering plants, goldenrod (Solidago Linnaeus spp. (Asterales: Asteraceae)) and mugwort (Artemisia vulgaris Linnaeus (Asterales: Asteraceae)), which should place them in prime locations for capturing flying pollinators.

Sexual selection on male body size, genital length and heterozygosity: Consistency across habitats and social settings.

Spatial and temporal variation in environmental factors and the social setting can help to maintain genetic variation in sexually selected traits if it affects the strength of directional selection. A key social parameter which affects the intensity of, and sometimes predicts the response to, mating competition is the operational sex ratio (OSR; ratio of receptive males to females). How the OSR affects selection for specific male traits is poorly understood. It is also unclear how sexual selection is affected by interactions between the OSR and environmental factors, such as habitat complexity, that alter key male-female interactions such as mate encounter rates. Here, we experimentally manipulated the OSR and habitat complexity and quantified sexual selection on male mosquitofish (Gambusia holbrooki) by directly measuring male reproductive success (i.e. paternity). We show that despite a more equitable sharing of paternity (i.e. higher levels of multiple paternity) under a male-biased OSR, selection on focal male traits was unaffected by the OSR or habitat complexity. Instead, sexual selection consistently, and significantly, favoured smaller bodied males, males with higher genome wide heterozygosity (based on >3,000 SNP markers) and males with a relatively long gonopodium (intromittent organ). Our results show that sexual selection on male body size, relative genital size and heterozygosity in this system is consistent across environments that vary in ecological parameters that are expected to influence mate encounter rates.

Sex in murky waters: algal-induced turbidity increases sexual selection in pipefish

Algal-induced turbidity has been shown to alter several important aspects of reproduction and sexual selection. However, while turbidity has been shown to negatively affect reproduction and sexually selected traits in some species, it may instead enhance reproductive success in others, implying that the impact of eutrophication is far more complex than originally believed. In this study, we aimed to provide more insight into these inconsistent findings. We used molecular tools to investigate the impact of algal turbidity on reproductive success and sexual selection on males in controlled laboratory experiments, allowing mate choice, mating competition, and mate encounter rates to affect reproduction. As study species, we used the broad-nosed pipefish, Syngnathus typhle, a species practicing male pregnancy and where we have previously shown that male mate choice is impaired by turbidity. Here, turbidity instead enhanced sexual selection on male size and mating success as well as reproductive success. Effects from mating competition and mate encounter rates may thus override effects from mate choice based on visual cues, producing an overall stronger sexual selection in turbid waters. Hence, seemingly inconsistent effects of turbidity on sexual selection may depend on which mechanisms of sexual selection that have been under study.Significance statementAlgal blooms are becoming increasingly more common due to eutrophication of freshwater and marine environments. The high density of algae lowers water transparency and reduces the possibility for fish and other aquatic animals to perform behaviors dependent on vision. We have previously shown that pipefish are unable to select the best partner in mate choice trials when water transparency was reduced. However, fish might use other senses than vision to compensate for the reduction in water transparency. In this study, we found that when fish were allowed to freely interact, thereby allowing competition between partners and direct contact between the fish, the best partner was indeed chosen. Hence, the negative effects of reduced water visibility due to algal blooms may be counteracted by the use of other senses in fish.

Male mate choice and insemination success under simultaneous versus sequential choice conditions

Theory predicts that males should be choosier when encountering potential mates simultaneously rather than sequentially because there is no opportunity cost. Consequently, when mate encounter rates vary across space and time males might benefit from plasticity in mate preferences to match prevailing social conditions, preferring high-quality mates when females are encountered frequently and showing no preferences when females are encountered rarely. Here we investigated how encounter type (i.e. simultaneous or sequential) alters male mate preferences for female size in the mosquitofish, Gambusia holbrooki . We found that male mosquitofish attempted to mate with a relatively large female significantly more often than a relatively small female when presented with two females simultaneously. In contrast, males showed no such preference when sequentially presented with two females. Further, males attempted more copulations with absolutely larger females irrespective of encounter type. Despite these behavioural patterns, however, neither male insemination success nor the number of sperm transferred was influenced by female size or the encounter type. Our results provide support for the prediction that male mate choice is stronger during simultaneous choice encounters, but suggest that insemination success in G. holbrooki is partly under female control.

Male-biased sex ratios and plasticity in post-insemination behaviour in the New Zealand stick insect Micrarchus hystriculeus

The duration of mate guarding by males is predicted to vary in accordance with the risk of sperm competition or mate encounter rate. Mate guarding is predicted to be prolonged under a male-biased sex ratio because the risk of sperm competition is high or the mate encounter rate is low. A consistently male-biased sex ratio should thus select for greater mate fidelity, and reduced plasticity in guarding behaviour, by males. Micrarchus hystriculeus Westwood (Phasmatodea) is a sexually size dimorphic stick insect in which males form prolonged post-insemination associations with their mates and whose populations have a consistently male-biased sex ratio. My laboratory experiments showed, as predicted, little plasticity by males in the duration of their post-insemination association. Although mate guarding duration is similar under male- and female-biased sex ratios, males clasped the genitalia of their mates significantly more often under a male-biased sex ratio, suggesting that males intensify their guarding with increasing risk of sperm competition.

Exploring simultaneous allocation to mating effort, sperm production, and body growth in male guppies

In several species, males increase their mating and sperm investment in the presence of unfamiliar females, the so-called Coolidge effect. Such an elevated reproductive effort is expected to be associated with a decreased investment in other costly traits, such as somatic growth and maintenance. How precopulatory, postcopulatory, and somatic investments interact one with each other, however, has been rarely evaluated simultaneously. We used the guppy (Poecilia reticulata), a polyandrous livebearing fish with alternative male mating tactics, to compare these 3 investments between males whose encounter rate with unfamiliar females was experimentally maintained at either high (high mate encounter rate [HER]) or low rate (low mate encounter rate [LER]) for 4 months. At the end of this period, HER males showed an increased sperm production as compared with their LER counterparts. This increment was accompanied with a reduction in the time spent following the female and by a shift from costly courtship displays to gonopodial thrusting, a less expensive coercive mating tactic. This effect may indicate a trade-off between these 2 components of male reproductive investment or may reflect a change in male optimal reproductive strategy associated with different mate encounter rate. In contrast, body growth was positively correlated with the increase in sperm production, and did not differ between experimental groups. Collectively, these results suggest that males did not vary their overall reproductive investment in response to their encounter rate with unfamiliar females, but differed significantly in the amount of resources available individually.

Anomalous diffusion and multifractality enhance mating encounters in the ocean

For millimeter-scale aquatic crustaceans such as copepods, ensuring reproductive success is a challenge as potential mates are often separated by hundreds of body lengths in a 3D environment. At the evolutionary scale, this led to the development of remote sensing abilities and behavioral strategies to locate, to track, and to capture a mate. Chemoreception plays a crucial role in increasing mate encounter rates through pheromone clouds and pheromone trails that can be followed over many body lengths. Empirical evidence of trail following behavior is, however, limited to laboratory experiments conducted in still water. An important open question concerns what happens in the turbulent waters of the surface ocean. We propose that copepods experience, and hence react to, a bulk-phase water pheromone concentration. Here we investigate the mating behavior of two key copepod species, Temora longicornis and Eurytemora affinis, to assess the role of background pheromone concentration and the relative roles played by males and females in mating encounters. We find that both males and females react to background pheromone concentration and exhibit both innate and acquired components in their mating strategies. The emerging swimming behaviors have stochastic properties that depend on pheromone concentration, sex, and species, are related to the level of reproductive experience of the individual tested, and significantly diverge from both the Lévy and Brownian models identified in predators searching for low- and high-density prey. Our results are consistent with an adaptation to increase mate encounter rates and hence to optimize reproductive fitness and success.

Frequency of Multiple Paternity in Gummy Shark, Mustelus antarcticus, and Rig, Mustelus lenticulatus, and the Implications of Mate Encounter Rate, Postcopulatory Influences, and Reproductive Mode

Elasmobranchs (sharks and rays) show an amazing diversity of reproductive modes and behaviors. Multiple paternity (MP) has been identified in all species where more than 1 litter has been investigated; yet neither direct nor indirect benefits from MP have been determined in elasmobranchs. This has led to the suggestion that MP in this group may simply be a product of convenience polyandry with variation in the frequency of MP driven by differences in mate encounter rates. Here, we use molecular markers to investigate polyandry and MP in 2 closely related and commercially important species of shark, Mustelus antarcticus and Mustelus lenticulatus. In total, 328 M. antarcticus embryos originating from 29 different mothers and 75 M. lenticulatus embryos originating from 19 different mothers were genotyped using 8 microsatellite loci. We find that MP occurs in both species. However, in both species, the majority of litters were sired by a single father. Our results do not support increased fecundity per se as a driver of MP. Further, our results do not suggest that high population densities with resulting high mate encounter rates generated by breeding aggregations necessarily lead to high frequencies of MP. Importantly, we note evidence of reproductive skew within polyandrous litters, which is a predicted outcome of postcopulatory mechanisms.

Molecular Evidence for Multiple Paternity in a Population of the Viviparous Tule Perch Hysterocarpus traski

Population density might be an important variable in determining the degree of multiple paternity. In a previous study, a high level of multiple paternity was detected in the shiner perch Cymatogaster aggregata, a species with high population density and a high mate encounter rate. The tule perch Hysterocarpus traski is phylogenetically closely related to C. aggregata, but it has relatively lower population density, which may result in distinct patterns of multiple paternity in these 2 species. To test the hypothesis that mate encounter rate may affect the rate of successful mating, we used polymorphic microsatellite markers to identify multiple paternity in the progeny arrays of 12 pregnant females from a natural population of tule perch. Multiple paternity was detected in 11 (92%) of the 12 broods. The number of sires per brood ranged from 1 to 4 (mean 2.5) but with no correlation between sire number and brood size. Although the brood size of tule perch is considerably larger than that of shiner perch (40.7 vs. 12.9, respectively), the average number of sires per brood in tule perch is much lower than that in shiner perch (2.5 vs. 4.6, respectively). These results are consistent with the hypothesis that mate encounter rate is an important factor affecting multiple mating.

Low Mate Encounter Rate Increases Male Risk Taking in a Sexually Cannibalistic Praying Mantis

Male praying mantises are forced into the ultimate trade-off of mating versus complete loss of future reproduction if they fall prey to a female. The balance of this trade-off will depend both on (1) the level of predatory risk imposed by females and (2) the frequency of mating opportunities for males. We report the results of a set of experiments that examine the effects of these two variables on male risk-taking behavior and the frequency of sexual cannibalism in the praying mantis Tenodera sinensis. We experimentally altered the rate at which males encountered females and measured male approach and courtship behavior under conditions of high and low risk of being attacked by females. We show that male risk taking depends on prior access to females. Males with restricted access to females showed greater risk-taking behavior. When males were given daily female encounters, they responded to greater female-imposed risk by slowing their rate of approach and remained a greater distance from a potential mate. In contrast, males without recent access to mates were greater risk-takers; they approached females more rapidly and to closer proximity, regardless of risk. In a second experiment, we altered male encounter rate with females and measured rates of sexual cannibalism when paired with hungry or well-fed females. Greater risk-taking behavior by males with low mate encounter rates resulted in high rates of sexual cannibalism when these males were paired with hungry females.

Population density and group size effects on reproductive behavior in a simultaneous hermaphrodite

BackgroundDespite growing evidence that population dynamic processes can have substantial effects on mating system evolution, little is known about their effect on mating rates in simultaneous hermaphrodites. According to theory, mating rate is expected to increase with mate availability because mating activity is primarily controlled by the male sexual function. A different scenario appears plausible in the hermaphroditic opisthobranch Chelidonura sandrana. Here, field mating rates are close to the female fitness optimum, suggesting that mating activity remains unresponsive to variation in mate availability.ResultsApplying an experimental design that aims at independent experimental manipulation of density and social group size, we find substantial increases in mate encounter rate with both factors, but no statistically detectable effects on mating rate in C. sandrana. Instead, mating rate remained close to the earlier determined female fitness optimum.ConclusionsWe demonstrate that mating rate in C. sandrana is largely unresponsive to variation in mate availability and is maintained close to the female fitness optimum. These findings challenge the prevailing notion of male driven mating rates in simultaneous hermaphrodites and call for complementary investigations of mating rate effects on fitness through the male sexual function.

Multiple mating and its relationship to brood size in pregnant fishes versus pregnant mammals and other viviparous vertebrates

We summarize the literature on rates of multiple paternity and sire numbers per clutch in viviparous fishes vs. mammals, two vertebrate groups in which pregnancy is common but entails very different numbers of embryos (for species surveyed, piscine broods averaged >10-fold larger than mammalian litters). As deduced from genetic parentage analyses, multiple mating by the pregnant sex proved to be common in assayed species but averaged significantly higher in fish than mammals. However, within either of these groups we found no significant correlations between brood size and genetically deduced incidence of multiple mating by females. Overall, these findings offer little support for the hypothesis that clutch size in pregnant species predicts the outcome of selection for multiple mating by brooders. Instead, whatever factors promote multiple mating by members of the gestating sex seem to do so in surprisingly similar ways in live-bearing vertebrates otherwise as different as fish and mammals. Similar conclusions emerged when we extended the survey to viviparous amphibians and reptiles. One notion consistent with these empirical observations is that although several fitness benefits probably accrue from multiple mating, logistical constraints on mate-encounter rates routinely truncate multiple mating far below levels that otherwise could be accommodated, especially in species with larger broods. We develop this concept into a "logistical constraint hypothesis" that may help to explain these mating outcomes in viviparous vertebrates. Under the logistical constraint hypothesis, propensities for multiple mating in each species register a balance between near-universal fitness benefits from multiple mating and species-idiosyncratic logistical limits on polygamy.

The interval between sexual encounters affects male courtship tactics in a desert-dwelling fish

Courtship displays are often important in determining male mating success but can also be costly. Thus, instead of courting females indiscriminately, males might be expected to adjust their signalling effort strategically. Theory, however, predicts that such adjustments should depend on the rate with which males encounter females, a prediction that has been subject to very little empirical testing. Here, we investigate the effects of female encounter rate on male courtship intensity by manipulating the time interval between sequential presentations of large (high quality) and small (low quality) females in a fish, the Australian desert goby Chlamydogobius eremius. Males that were presented with a small female immediately after a large female reduced their courtship intensity significantly. However, males courted large and small females with equal intensity if the interval between the sequential presentations was longer. Our results suggest that mate encounter rate is an important factor shaping male reproductive decisions and, consequently, the evolutionary potential of sexual selection.

ALLEE EFFECT LIMITS COLONIZATION SUCCESS OF SEXUALLY REPRODUCING ZOOPLANKTON

Understanding the dynamics of populations at low density and the role of Allee effects is a priority due to concern about the decline of rare species and interest in colonization/invasion dynamics. Despite well-developed theory and observational support, experimental examinations of the Allee effect in natural systems are rare, partly because of logistical difficulties associated with experiments at low population density. We took advantage of fish introduction and removal in alpine lakes to experimentally test for the Allee effect at the whole-ecosystem scale. The large copepod Hesperodiaptomus shoshone is often extirpated from the water column by fish and sometimes fails to recover following fish disappearance, despite the presence of a long-lived egg bank. Population growth rate of this dioecious species may be limited by mate encounter rate, such that below some critical density a colonizing population will fail to establish. We conducted a multi-lake experiment in which H. shoshone was stocked at densities that bracketed our hypothesized critical density of 0.5-5 copoepods/m3. Successful recovery by the copepod was observed only in the lake with the highest initial density (3 copepods/m3). Copepods stocked into small cages at 3000 copepods/m3 survived and reproduced at rates comparable to natural populations, confirming that the lakes were suitable habitat for this species. In support of mate limitation as the mechanism underlying recovery failure, we found a significant positive relationship between mating success and density across experimental and natural H. shoshone populations. Furthermore, a mesocosm experiment provided evidence of increased per capita population growth rate with increasing population density in another diaptomid species, Skistodiaptomus pallidus. Together, these lines of evidence support the importance of the Allee effect to population recovery of H. shoshone in the Sierra Nevada, and to diaptomid copepods in general.

Heterospecific mating and partial prezygotic reproductive isolation in the planktonic marine copepods Centropages typicus and Centropages hamatus

Using three‐dimensional (3D) video observations in laboratory experiments, I describe interspecific and intergeneric mating behaviors and motility patterns of the common planktonic marine copepods Centropages typicus, Centropages hamatus, and Temora longicornis. These observations are then used to estimate heterospecific and conspecific male mate‐search volume rates and mate encounter rates in North Sea Centropages populations. Behavioral prezygotic reproductive isolation between Centropages species is incomplete, since males of each species pursued, contacted, captured, and, in rare cases, placed a spermatophore on the urosome of heterospecific females. T. longicornis males also detected the diffusible pheromone trail and pursued C. typicus females to the point of mate contact. Male mate‐search tracking behavior was equally effective on diffusible pheromone trails of heterospecific and conspecific females, indicating that pheromone and hydromechanical precontact mating cues lack species specificity. Males attempted mating with both heterospecific and conspecific females at high frequencies in the laboratory, and species recognition is inferred to occur through contact chemosensory cues or morphological incompatibility between species. Heterospecific mate encounter rates in the North Sea were maximal in late summer (August), up to~2,000 encounters m−3 d−1, and were comparable in magnitude to conspecific encounter rates during the same period. C. typicus females experience the highest incidence of heterospecific mating interactions, since they encounter heterospecific males at rates up to 100+ encounters female−1 d−1, ca. one order of magnitude higher than encounter rates with conspecifics. Heterospecific mating attempts may be a common feature of the reproductive ecology of planktonic copepods and may incur substantial fitness costs to the individuals involved.