Sperm competition. II-post-copulatory guarding

A two round sperm competition model is analysed to determine which male strategy is advantageous for fertilization of a given set of eggs; guarding a particular female or searching for another copulation. A guarding male is one who would guard if he mates in the first round (which may not occur) whilst a non-guarding male decides on how much sperm to allocate if given the opportunity to inseminate a female in round one. Guarding behaviour is defined in terms of a probability of preventing a further insemination if challenged by a rival male. Sperm success with a single female obeys the "raffle principle". An evolutionarily stable strategy (ESS) approach is used to ascertain the best non-guarding ejaculation strategy. We show that for each fixed proportion of guarders in the population the strategies are ordered and that only a single guarding strategy need be considered. The model predicts that there will be evolution to either the guarding strategy or a single non-guarding strategy or a polymorphic combination of guarding and some (or all) of the non-guarding strategies. The conditions for coexistence to occur were shown to be rare in comparison to those necessary for a monomorphism. Copyright 1999 Academic Press.     

Compromised strategy resolves intersexual conflict over pre-copulatory guarding duration

Summary An evolutionarily stable strategy (ESS) on pre-copulatory mate-guarding duration is separately obtained for males and females, by assuming that either the male or female can control perfectly the timing of guarding. A difference between sexes in an ESS brings on an intersexual conflict, in particular when the ESS of the actively searching sex (usually male) is longer than that of the other. We analyse two extreme situations, in which the female mating stages are either perfectly synchronized or uniformly distributed. The analysis reveals that (1) the male ESS for guarding duration is longer than the female ESS in the synchronized case if the sex ratio is male-biased, (2) the difference in ESSs is higher for a more male-biased sex ratio, less guarding costs or a higher encounter rate, and (3) an asynchronous female mating cycle extends the conflict region towards female-biased sex ratios. We show by including conflict costs in fitnesses of both sexes that intersexual conflict may be resolved by a compromised solution, where the starting time of mate guarding is an intermediate value between the ESSs of the two sexes. This compromised strategy depends on both fitness increments of winning the conflict and physical power in controlling the opponent and tends to approach the ESS of the commoner sex in highly biased sex ratios. If both actors engaged in a conflict have enough information on each other, a compromise without an overt struggle may be reached.     

Models on butterfly protandry: Virgin females are at risk to die

Current models on protandry in butterflies assume that females are mated instantaneously upon eclosion. However, for most butterfly species this assumption is not realistic. In this paper a model is formulated in which the mating rate depends on both male and female density. Given the female presence curve, protandry is an evolutionarily stable strategy (ESS) for males. The evolutionarily stable amount of protandry decreases with increasing death rate and decreasing encounter rate. Given the male presence curve, protandry also is an ESS for females. However, male and female ESS are not identical; moreover, in the present model a simultaneous ESS does not exist. Protandry critically depends on the assumption that females mate only once, whereas males are capable of multiple mating. If females too are capable of multiple mating, absence of protandry is the ESS for males as well as females. The model predicts that protandry depends on population density: protandry should be more pronounced in populations with high density than in populations with low density. Protandry also depends on sex ratio. It becomes more pronounced when the proportion of males among emerging adults increases.     

The male's dilemma: Increased offspring production is more paternity to steal

Summary Large potential effects of male care on the number of offspring females successfully raise are not sufficient to select for caring males because of the pervasive importance of mating competition. Males face a version of the social dilemma, in which increased production increases the pay-off for theft. Models of the allocation of male effort partitioned between caring for babies and competing for paternity show that the optimal allocation to care is very low under a wide range of conditions. Like sex allocation where the alternatives are male versus female function or sons versus daughters, the pay-offs to one alternative are always strongly frequency dependent. Because that alternative (male function, sons, male mating effort) pays so well when rare, it cannot remain rare under most conditions. Here we consider the consequences of partitioning mating effort into mate guarding and all other forms of mating conflict. If a male gets all his partner's conceptions while guarding, gaining them at a constant rate, there are two possible regions of stability. The evolutionarily stable strategy (ESS) depends on a parameter scaling the decisiveness of (non-guarding) mating conflict. When marginal returns from conflict decrease with scale, almost all effort goes into guarding. When marginal returns increase, the ESS devotes all effort to mating. Even when the potential effect of care is large, male equilibrium strategies allocate little effort to it. We also report the results of computer simulations showing that care increases if gains from guarding saturate quickly, so that a male is assured of the paternity of most of his partner's offspring with little guarding, and consequently the pool of unguarded conceptions open to competion shrinks sharply. But even when the male's dilemma is very much reduced, it still substantially limits the allocation to care. The results of both computer simulations and mathematical analysis converge with other lines of evidence that mating has much stronger effects than parenting in shaping male strategies.     

The Comparative Biology of Genetic Variation for Conditional Sex Ratio Behavior in a Parasitic Wasp, Nasonia Vitripennis

Using genetic markers, we tracked the sex ratio behavior of individual females of the parasitic wasp, Nasonia vitripennis, in foundress groups of size 1, 2, 4, 8 and 16. Comparison of 12 isofemale strains extracted from a natural population reveals significant between-strain heterogeneity of sex ratios produced in all sizes of foundress group. Under simple assumptions about population structure, this heterogeneity results in heterogeneity of fitnesses. The strains differ in their conditional sex ratio behavior (the sex ratio response of a female to foundress groups of different sizes). Females of some strains produce more males as foundress group size increases (up to size eight). Females of another strain produce more males when not alone but do not respond differentially to group size otherwise. Females of two other strains show no conditional sex ratio behavior. Females of only two strains behave differently in foundress groups of size 8 and 16. Correlation and regression analyses indicate that the strains differ significantly in their fit to the predictions of an evolutionarily stable strategy (ESS) model of conditional sex ratio behavior. Such heterogeneity contradicts the notion that females of this species possess conditional sex ratio behavior that is optimal in the ESS sense. The results imply that this ESS model is useful but not sufficient for understanding the causal basis of the evolution of this behavior in this species. This is the first report on the sex ratio behavior of individual females in multiple foundress groups in any species of parasitic wasp. Data of this type (and not foundress group or ``patch'''' sex ratios) are essential for testing evolutionary models that predict the sex ratio behaviors of individuals. We suggest that a test for an ESS model include the answers to two important questions: 1) is the model quantitatively accurate? and 2) is there reasonable evidence to indicate that natural selection has caused individuals to manifest the ESS behavior?     

Male Mating Tactics in the Shrimp Palaemonetes pugio (Decapoda, Caridea): Precopulatory Mate Guarding vs. Pure Searching

The guarding of females approaching a limited period of sexual receptivity is a common mating tactic of males. In many decapod crustaceans, such as the shrimp Palaemonetes pugio , females can only copulate during a short period after a reproductive molt. It has been predicted that mate guarding by males (pre-copula) evolves in such species if sex ratios are not highly female-biased and if males can detect the molt stage of the female. The mating tactics of males were investigated in P. pugio . Time-lapse video observations were made on interactions among two males, a pre-molt female, and an inter-molt female (20 replicates). There was no evidence that males recognized a pre-molt female until 24 h before its molt. Significant numbers of male contacts with pre-molt females occurred 1 h before and after the female molt. Copulation took place within 1–3 min of the molt. No behavior commonly associated with mate guarding in decapods was observed – no clasping, agonistic behavior, or close association. It is concluded that the male's mating tactic is pure searching, wherein males haphazardly contact many females in order to find a receptive one. The high encounter rate in nature of these very mobile, aggregated shrimps is proposed as the factor responsible for the evolution of pure searching. It is hypothesized that pure searching is the male tactic of the many species of decapod shrimps with small males, sexually monomorphic cheliped weapons, and aggregated populations.     

Female Reproductive Cycle and Sexual Conflict over Precopulatory Mate-guarding in Thermosphaeroma (Crustacea, Isopoda)

In species with time-limited opportunities for insemination, precopulatory mate-guarding is expected to coevolve with the duration of female reproductive cycles. Despite this adaptation to female characteristics, it may also be advantageous for males to adjust the duration of guarding with respect to sex ratio because the benefits of guarding are dependent on the availability of females. If female fitness is reduced because of guarding, male guarding behavior leads to intersexual conflict. We studied these aspects of male mate-guarding behavior in two closely related, thermal-spring isopods (Thermosphaeroma). First, guarding duration showed species specificity which was related to the duration of reproductive cycle; cycle length for females and duration of guarding by males in T. milleri were twice as long as in T. thermophilum. Second, males in both species adjusted their guarding duration with sex ratio, guarding longer when a competing male was present. Third, in T. thermophilum, ovarian development began immediately after the birth of the previous brood and continued through guarding, sexual molt and post-molt periods until oviposition, whereas in T. milleri, ovarian development was largely postponed until the post-molt period. Because guarding during ovary provisioning periods may be costly for females, we tested the existence of intersexual conflict over guarding duration in T. thermophilum. We compared the duration of guarding of control pairs with those of pairs in which either male guarding ability or female ability to resist guarding was reduced experimentally. Guarding durations for manipulated and control males were equal, but manipulated females were guarded longer, suggesting that conflict exists and that females can effectively shorten guarding duration by their behavior. Moreover, we suggest that selection in the context of intersexual conflict may play an important role in the evolution of delayed oviposition and sperm-storage organs in mate-guarding crustaceans.     

Quantification of sexual dimorphism in Asellus aquaticus (Crustacea: Isopoda) using outline approaches

A marked sexual dimorphism is often observed in arthropods species in which males perform precopulatory mate guarding. It is generally thought to reflect the influence of sexual selection. Until now, sexual dimorphisms associated with mate guarding have mainly been qualitatively described. However, assessing the effects of sexual selection on sexual dimorphims requires a preliminary quantitative assessment of differences in morphology between sexes. Using Fourier analyses, we tested if morphological dimorphisms could be quantitatively assessed in the isopod Asellus aquaticus . In addition, we checked whether sexual dimorphism in shape was exclusively related to mate guarding through considering characters that are not, a priori , implicated in mating behaviour. To assess the potential role of sexual selection in shaping morphology, we then examined how dimorphic characters could influence males' pairing success. Three characters (pleotelson, paraeopods 4 and 5) differed significantly in shape between males and females. In addition, two characters (pleotelson and paraeopods 4) differed in shape between guarding males and non-guarding males, with the latter being closer in shape to females. This suggests that sexual selection may be partly responsible for the observed morphological divergence between sexes in A. aquaticus .  © 2002 The Linnean Society of London, Biological Society of the Linnean Society , 2002, 77 , 523–533.     

The Conflict between Nest Guarding and Mate Guarding in Penduline Tits (Remiz pendulinus)

In penduline tits (Remiz pendulinus), polygynous males build several very elaborate nests successively during one breeding season to attract females. The time and effort invested in nest building is positively related to their mating success. Intraspecific competition for nest material resulting in nest material theft, delays males' nest building progress which in turn decreases their mating success. Therefore, a nest guarding strategy was predicted. In many bird species, males develop some kind of paternity strategy during the female fertile phase. However, as nest building and the female fertile phase frequently coincide, one would expect a trade-off between these strategies. In this study we determined in particular how nest guarding in males conflicts with their mate guarding behaviour in penduline tits. Our results show that nest building is costly in terms of a male's time budget. Thieves benefit by increasing their rate of acquiring nest material which reduces the effort invested in nest building. Nest guarding is an efficient strategy to avoid thieves, as indicated by the negative relation between the presence of the male near the nest and the frequency of nest material theft. Nest guarding is required for the whole building period but males, however, increased their mate guarding effort during the peak fertile phase to ensure their paternity. The data suggest that, for the trade-off “mate guarding versus nest guarding”, paternity insurance seems to be more important.     

Parental care and social organization of the spiny eel, Aethiomastacembelus platysoma, in Lake Tanganyika

This is the first report demonstrating the occurrence of parental care in mastacembelids. Social organization and parental care of a spiny eel Aethiomastacembelus platysoma were studied in Lake Tanganyika. Both males and females maintained individual territories though the frequency of aggressive interaction was low. The male guarded offspring in a rock hole within its territory. The egg size was large (2.5–2.7 mm in diameter) and the brood size in a nest was 5.7 on average in spite of more oocytes in the ovary (65 large oocytes on average). The duration of guarding was around 30 days after hatching and the young became independent just after they began to feed. Guarding males seldom attacked fishes that approached the nest, and often went out of the nest to forage though the stomach contents of guarding males were less than those of non-guarding males. Compared with Tanganyikan cichlid fishes that show prolonged parental care at open sites, the post-hatching guarding interval is short and the egg size is large, which seem to be traits common to fishes that utilize closed spaces as guarding sites in the lake.     

Part-Time Mate Guarding Affects Paternity in Male Reed Buntings (Emberiza schoeniclus)

Male mate guarding by close following has been reported in many socially monogamous bird species and is generally believed to function as a paternity guard. Many aspects of the dynamics and effectiveness of this behavior are still however poorly understood. Here, we describe the temporal variation in mate guarding behavior in male reed buntings (Emberiza schoeniclus) with a particular focus on how males allocate their mating effort between mate guarding and extrapair mating in a context of intense sperm competition. In our highly synchronous study population most males have to balance the simultaneous and mutually exclusive demands of mate guarding and seeking extrapair copulations (EPCs). We found that males frequently switched between guarding their mates and performing intrusions to neighboring territories. Both activities seemed to have significant fitness payoffs, as male mate guarding effort had a positive effect on paternity, and a large fraction of extrapair fertilizations occurred during the days when the sire guarded its own female. The reed bunting is thus an example of how discontinuous or part‐time mate guarding can still be effective in securing paternity. Female reed buntings were not particularly active in initiating EPCs as they never were observed performing extraterritorial forays. We argue that the absence of female‐initiated EPCs is a prerequisite for males to trade mate guarding against seeking EPCs. Otherwise, if females circumvent male mate guarding by timing their EPCs to periods of male absence, males should guard their mates almost continuously or rely on alternative paternity guards.     

Factors influencing mate guarding and territory defence in the stitchbird (hihi)

Socially monogamous male birds are predicted to maximise their reproductive success by pursuing extra-pair copulations (EPCs) while engaging in anti-cuckoldry behaviour such as mate guarding. In the stitchbird, Notiomystis cincta, high levels of forced EPCs and a high proportion of nestlings resulting from extra- pair fertilisations lead to the prediction that males of this species should exhibit intense paternity guarding behaviours. While studying an isolated stitchbird population on Tiritiri Matangi Island New Zealand (3636'S, 17453'E), I collected daily behavioural data throughout the breeding season from 15 males in 2000/01 and 27 males in 2001/02. In this study, male stitchbirds demonstrated clear paternity guarding by exhibiting: (1) an increased likelihood of being close to their mate during her fertile period, (2) an increased initiation of mate contact during her fertile period, (3) switching from site-specific territorial defence during the pre-fertile period to defending an area centring on the their female partners location during her fertile period, and (4) an increased following of the female to communal feeding sites outside the territory during her fertile period. For polygynous males, mate guarding and territorial defence were conditional on which of their females was fertile. Additional evidence supporting the hypothesis that mate guarding in this species is a form of paternity assurance, rather than protection from harassment, is that males protected their partner from harassment by other stitchbird males but did not intervene when females were harassed by male bellbirds, Anthornis melanura. While mate-guarding intensity in many species is conditional on the stage of female fertility, male stitchbirds also modified their behaviour depending on the location of the female and the rate of intrusions by extra-pair males. Resident males adopted a best-of-a-bad-job tactic when they were unable to locate their female by defending an area around her last known location. Furthermore, when the rate of intrusions by extra-pair males increased they traded-off the area they could defend within their territory against their ability to guard the female. Territory takeovers were uncommon, but when they did occur older males displaced younger males and healthy birds displaced sick ones. Contrary to the prevailing view that mate guarding is a male response to female infidelity, male stitchbirds appear to use mate guarding primarily to prevent paternity losses from forced EPCs. Future assessments of mate guarding function should consider the possibility that mate guarding involves a combination of conflict and co-operation between the sexes.     

The evolution of parental care in the context of sexual selection: a critical reassessment of parental investment theory

Males and females are often defined by differences in their energetic investment in gametes. In most sexual species, females produce few large ova, whereas males produce many tiny sperm. This difference in initial parental investment is presumed to exert a fundamental influence on sex differences in mating and parental behavior, resulting in a taxonomic bias toward parental care in females and away from parental care in males. In this article, we reexamine the logic of this argument as well as the evolutionarily stable strategy (ESS) theory often used to substantiate it. We show that the classic ESS model, which contrasts parental care with offspring desertion, violates the necessary relationship between mean male and female fitness. When the constraint of equal male and female mean fitness is correctly incorporated into the ESS model, its results are congruent with those of evolutionary genetic theory for the evolution of genes with direct and indirect effects. Male parental care evolves whenever half the magnitude of the indirect effect of paternal care on offspring viability exceeds the direct effect of additional mating success gained by desertion. When the converse is true, desertion invades and spreads. In the absence of a genetic correlation between the sexes, the evolution of paternal care is independent of maternal care. Theories based on sex differences in gametic investment make no such specific predictions. We discuss whether inferences about the evolution of sex differences in parental care can hold if the ESS theory on which they are based contains internal contradictions.     

Changes in male physiological condition during brooding activities in a natural population of a stream goby, <Emphasis Type="Italic">Rhinogobius</Emphasis> sp.

We investigated the change in physical condition of males during paternal care in a natural population of a stream goby Rhinogobius sp. LD. Hepatosomatic index (HSI) and Condition factor (K) of males guarding an egg mass were higher than those of non-breeding males. In the first brooding cycle, the HSI of males decreased with the progress of embryo development. Moreover, in the second brooding cycle, both the HSI and K decreased, suggesting that the energy reserves in the somatic tissue were used when the liver reserves were exhausted as a source of energy for paternal activities. After paternal care, males showed reduced K and were unable to recover HSI immediately. Comparisons of the physical condition between non-guarding males after the first brooding cycle and males guarding an early stage clutch in the second brooding cycle indicated that males which could recover HSI and K were able to have a second brooding cycle. This suggested that the physical condition determines participation in breeding activity and the number of brooding cycles, greatly influencing male reproductive success.     

Age-Related Variation in Mate-Guarding Intensity in the Bluethroat (Luscinia s. svecica)

Female extra‐pair copulations (EPCs) have selected for male paternity guarding strategies in many bird species. In the bluethroat, Luscinia s. svecica, males guard their mates closely during the last 2 d before the start of egg laying, but there is great individual variation in the intensity of mate guarding. Here we show that some of this variation is related to male age. Old males guarded their mates with much lower intensity and sang more than young males, although the latter difference was not statistically significant. Controlling for male age, male and female coloration and size were not significantly related to the intensity of mate guarding. We have previously shown that young and old males had a similar paternity loss in their own broods. On the other hand, old males were far more successful than young males in achieving extra‐pair fertilizations. These patterns suggest that young and old males have different trade‐offs between preventing paternity loss in own nest and gaining paternity in others, because male skills in obtaining EPCs improve with experience and/or because of female preferences for old males as copulation partners. There were no significant relationships between paternity and male mate‐guarding behaviour during the fertile period, indicating that mate guarding is not a very effective paternity‐assurance strategy in the bluethroat.     

Parental investment of male two-spotted goby, Gobiusculus flavescens (Fabricius)

The trade-off between parental care and feeding was studied in the male two-spotted goby (Gobiusculus flavescens F.). Two temperatures, 8.5 degrees C and 13.0 degrees C, were used, with five replicates at each temperature, in order to determine whether temperature influenced parental behaviour. In each replicate, two males and four females were introduced to an aquarium, where the males chose between two nests and courted the females. In each replicate, one male spawned. After spawning, the males guarded the eggs until hatching. The guarding males' behaviour was recorded with a video camera twice a day (15 min each time), once before and once after they were fed. The male's condition (c-factor) was calculated at the start of the experiment and after egg hatching. The eggs were spawned in an artificial nest (half of a PVC tube), and attached to the nest in a single layer. The areas with eggs (representing brood size) were marked after spawning and the fry counted after hatching (which was used to calculate area hatched). Numbers of prey eaten (plankton) and number of aggressive encounters between the guarding male and the other fishes were recorded. Time spent in the nest and time used on fertilisation, fanning and cleaning were estimated and related to egg age, brood size, hatching success, temperature and food availability (no food or food).The results showed that feeding (expected to influence future reproduction) decreased and parental expenditure (current reproduction) increased, as the eggs developed (became closer to independence). Parental expenditure was significantly higher at 13.0 degrees C than at 8.5 degrees C, presumably due to higher oxygen demands by the eggs, and a greater risk of egg-infections. The c-factor of the males guarding eggs decreased over time, in contrast to the non-guarding males' c-factor. Guarding males' aggressiveness decreased as the eggs got older, but increased just before hatching. A possible explanation for this could be the decreasing intrusion by the non-guarding male and females caused by high aggressive behaviour by the guarding male early in the brood cycle. The exploitation of the nest (percentage of total nest area covered by eggs) seemed to determine the amount of parental expenditure and loss of condition, while brood size (area of eggs) had no effect.     

Mated pairs of owl monkeys (Aotus nancymaae) exhibit sex differences in response to unfamiliar male and female conspecifics

In socially monogamous species, mate‐guarding could be a reproductive strategy that benefits both males and females, especially when males contribute to parental care. By actively guarding mates, males may reduce their chances of being cuckolded, whereas females that mate‐guard may reduce the likelihood that their mates will desert them or acquire additional mates, and hence limit or reduce paternal care of offspring. Owl monkeys (Aotus spp.) are socially monogamous with biparental care of young and, hence, potential beneficiaries of mate‐guarding. We presented mated pairs of captive owl monkeys (A. nancymaae) with unfamiliar male and female conspecifics, to determine if either member of the pair exhibits intraspecific aggression toward an intruder or stays close to its mate, behaviors indicative of mate‐guarding. Male mates were more responsible for the maintenance of close proximity between mates than females. Male mates also exhibited elevated levels of behavior that signify arousal when presented with a male conspecific. These responses by mated male owl monkeys are consistent with patterns that may help prevent cuckoldry. Am. J. Primatol. 72:942–950, 2010. © 2010 Wiley‐Liss, Inc.     

The adaptiveness of intense contact mate guarding by males of the emerald damselfly,Lestes sponsa (Odonata, Lestidae): The male’s perspective

We studied the mating system of the emerald damselflyLestes sponsa. All males showed intense contact mate guarding by holding the female in tendem during the entire oviposition period. Our findings support the predictions made by Alcock (1994) about the occurrence of intense mate guarding: (1) a high female receptivity after copulation, (2) a high male capacity to resist takeovers, (3) sperm precedence, (4) a high operational sex ratio, (5) a high male density, (6) high access by rivals to mated females, (7) low energy expenditure, (8) a low risk of guarding, and (9) a short interval between copula and oviposition. This indicates a positive cost-benefit balance for this behavior, at least in males. A comparison within the genusLestes suggests that the male-biased sex ratios and the ease with which mated females are detected have been strong selection pressures in the evolution of intense contact mate guarding.     

Sperm competition games: sperm selection by females

We analyse a co-evolutionary sexual conflict game, in which males compete for fertilizations (sperm competition) and females operate sperm selection against unfavourable ejaculates (cryptic female choice). For simplicity, each female mates with two males per reproductive event, and the competing ejaculates are of two types, favourable (having high viability or success) or unfavourable (where progeny are less successful). Over evolutionary time, females can increase their level of sperm selection (measured as the proportion of unfavourable sperm eliminated) by paying a fecundity cost. Males can regulate sperm allocations depending on whether they will be favoured or disfavoured, but increasing sperm allocation reduces their mating rate. The resolution of this game depends on whether males are equal, or unequal. Males could be equal: each is favoured with probability, p, reflecting the proportion of females in the population that favour his ejaculate (the 'random-roles' model); different males are favoured by different sets of females. Alternatively, males could be unequal: given males are perceived consistently by all females as two distinct types, favoured and disfavoured, where p is now the frequency of the favoured male type in the population (the 'constant-types' model). In both cases, the evolutionarily stable strategy (ESS) is for females initially to increase sperm selection from zero as the viability of offspring from unfavourable ejaculates falls below that of favourable ejaculates. But in the random-roles model, sperm selection decreases again towards zero as the unfavourable ejaculates become disastrous (i.e. as their progeny viability decreases towards zero). This occurs because males avoid expenditure in unfavourable matings, to conserve sperm for matings in the favoured role where their offspring have high viability, thus allowing females to relax sperm selection. If sperm selection is costly to females, ESS sperm selection is high across a region of intermediate viabilities. If it is uncostly, there is no ESS in this region unless sperm limitation (i.e. some eggs fail to be fertilized because sperm numbers are too low) is included into the model. In the constant-types model, no relaxation of sperm selection occurs at very low viabilities of disfavoured male progeny. If sperm selection is sufficiently costly, ESS sperm selection increases as progeny viability decreases down towards zero; but if it is uncostly, there is no ESS at the lowest viabilities, and unlike the random-roles model, this cannot be stabilized by including sperm limitation. Sperm allocations in the ESS regions differ between the two models. With random roles, males always allocate more sperm in the favoured role. With constant types, the male type that is favoured allocates less sperm than the disfavoured type. These results suggests that empiricists studying cryptic female choice and sperm allocation patterns need to determine whether sperm selection is applied differently, or consistently, on given males by different females in the same population.     

Chemically-mediated pre-mating behavior in two tetranychid species

We used bioassays to evaluate the arrestment response of male twospotted spider mite, Tetranychus urticae Koch, and Banks grass mite, Oligonychus pratensis (Banks) to whole-body extract from conspecific quiescent deutonymphs. We examined the effect of previous behavior on mite response to extract from female quiescent deutonymphs. We also examined male arrestment and guarding behavior in response to 2 extract concentrations and to extract from male quiescent deutonymphs. Male T. urticae and O. pratensis exhibited similar changes in their behavior in response to the different extracts with which we presented them. Males that were guarding quiescent deutonymphs immediately prior to testing spent more time in an untreated 3.5-mm-diameter circle than did males that were previously engaged in other behaviors. However, when nonguarding males were presented with extract of conspecific female quiescent deutonymphs they remained in the stimulus circle as long as guarding males did in an untreated circle. Arrestment duration of nonguarding males increased at 2 higher extract concentrations. The arrestment response was not exclusive to extracts from female quiescent deutonymphs; nonguarding male mites were also arrested by extracts from male quiescent deutonymphs, although for a shorter time. Duration of arrestment was related roughly to size differences between male and female quiescent deutonymphs. Furthermore, males did not show an exclusive preference for guarding conspecific male or female quiescent deutonymphs. It seems unlikely that the arrestant is a specific sex pheromone. Rather, male mites probably use the arrestant as a necessary cue to focus their attention on an individual that may be a suitable mate. Other tactile, visual, and chemical stimuli may then help males to decide whether to remain and assume mate guarding behaviors.