Indirect female choice mediated by sex pheromones in the hermit crab Pagurus filholi

Males of the hermit crab Pagurus filholi perform precopulatory guarding behavior, and solitary males often show aggressive behavior to take away guarded females. Males behave coercively while guarding females, so direct mate choice by females seems difficult in such a situation. By performing several experiments we examined possible indirect female choice of hermit crab. Males were attached to a shell by their left cheliped to look like guarding pairs (fake guarding pairs). The shells were filled with cotton containing either seawater or pheromone water. The fake guarding pair with only seawater caused male–male combat in 60% of trials whereas with pheromone water combats occurred in 88% of trials. Mean duration of male–male combat was significantly longer in trials with drops of seawater containing pheromones than in those without pheromones. These results suggest guarding pairs themselves cause male–male combat by visual stimulation, that female sex pheromones have further significant function in the recognition of guarding pairs and intensification of male–male combat, and that females release sex pheromones while they are guarded. As a result of the combat, the larger male ended up guarding a female. This strongly suggests that females choose males indirectly by exploiting male–male competition induced by sex pheromones under male coercive behavior.     

Social interactions associated with reproduction in the bushveld gerbil <Emphasis Type="Italic">Gerbilliscus leucogaster</Emphasis>

The study of social behaviour can give us important insights into the social and mating system of a species or population. We investigated male–female interactions in captive bushveld gerbils, Gerbilliscus leucogaster, to gain insight into the social behaviour and mating structure. We conducted two experiments. In experiment 1, we studied the interactions between two strangers for a week in a three-tank setup, which tested behavioural variation spatially and temporally. Although tolerance between strange males and females increased with time, it remained generally low throughout the experiment. Females appeared to be cautious of males initially, but they later became aggressive towards them. Males showed an increase in submissive behaviour over time. The ‘home’ cage did not appear to be defended by either sex. In experiment 2, we investigated the social interactions of male–female pairs during pregnancy and lactation. Aggression persisted throughout the study and amicable behaviour was low; females were much more aggressive than males. We did not detect changes in social behaviour with the progression of pregnancy and lactation. We suggest that pair bonding is unlikely and that promiscuity is the most probable mating system. Female aggression may be related to mate choice prior to mating and mate exclusion thereafter, while it may be a response to infanticide risk during pregnancy and lactation.     

Size-assortative pairing in the lotic amphipod <Emphasis Type="Italic">Gammarus zaddachi</Emphasis>, an examination of hypotheses and the influence of current speed

Three main hypotheses have been put forward to explain size-assortative pairing in gammarid amphipods: microhabitat separation, sexual selection and loading constraint. In order to determine which hypothesis best explains this phenomenon in the estuarine species Gammarus zaddachi, I first measured the body lengths and dry weights of precopula pairs collected from two field sites with substantially different current speeds. Second, I performed three laboratory experiments in order to estimate the importance of the following processes: (1) male choice; (2) male–male competition and (3) male–female acceptability. The loading constraint hypothesis seemed best supported by the data in that field-collected male G. zaddachi size correlated well with female size in precopula pairing in both fast and slow flowing water. In the laboratory, males preferred females of their same size group (large versus small), and ‘won’ them in the male–male competition experiments. Size-assortative pairing is thus likely a consequence of the loading constraints imposed upon these males by virtue of them having to carry and manoeuvre their partners through flowing water, while attempting to maintain station in an optimal microhabitat. Males may therefore forego the largest, most fecund females, in favour of a practicable payload (small male–large female pairings were rare). However, there seems to be a lower limit to this selection, indicated by the high degree of cannibalism on small females by large males.     

Role of Contests in the Scramble Competition Mating System of a Leaf Beetle

Male competition for mates can occur through contests or a scramble to locate females. We examined the significance of contests for mates in the leaf beetle Chrysomela aeneicollis, which experiences a short breeding season. During peak mating season, 18–52% of beetles are found in male-female pairs, and nearly half of these are copulating. Sex ratios do not differ from parity, females are larger than males, and positive size-assortative mating occurs. Males fight (2–4% of beetles) over access to females, and disruption of mating usually follows these contests. In the laboratory, we compared mating and fighting frequencies for males found in mating pairs (field-paired) and single males placed into an arena with a field-paired female. Mating pairs were switched in half of arenas (new male-female pairs) and maintained in the other half. For 2 days, each male was free to move about and fight; thereafter males were tethered to prevent contests. Mating frequencies were significantly greater for field-paired than single males in both situations. Male size was not related to mating frequency; however, large females received more matings than small ones. These data suggest that males fight for high quality females, but otherwise search for as many matings as possible.     

Mate choice and sexual conflict in <Emphasis Type="Italic">Brachionus plicatilis</Emphasis> (Rotifera)

Male mate choice in Brachionus plicatilis is based on information from coronal receptors and contact chemoreception of a glycoprotein signal on the body surface of females. Male mating behavior is modulated by mating signal quality and strength, which determines male mating persistence and circling intensity. We probe the sensory abilities of males by better resolving their discrimination of female age. Males preferentially initiate mating with young females, but not too young. Males circle 0.2 h old females just hatched from resting eggs only about 1/2 as frequently as 12 h old females. Males are even more discriminating of females with whom they copulate, preferring 3 h old females significantly more than 0.2 h or 6 h old females. In contrast, males cannot distinguish between virgin females and those who have already copulated. Similarly, males circled and copulated with females hatching from resting eggs with equal frequency as those hatching from amictic eggs. The counterpoint to male mate choice is female resistance to mating. When males contact females, the females respond with one of four behaviors: no response, acceleration, foot flipping, or coronal retraction. In 65% of male–female encounters, there was no initial response by females. However, when males began circling females, females accelerated 11.1 times more often than when males were absent. The second type of evasive female behavior was foot-flipping, which tends to knock off males attempting to circle the female. In the presence of circling males, females performed foot-flipping behavior three times more often than in the absence of males. Coronal retraction, where they stop swimming and withdraw their corona, was observed less frequently than acceleration or foot-flipping, and there was no difference in the presence or absence of males. These data are interpreted in the context of sexual conflict, where the behaviors that optimize male and female fitness differ. Guest editors: S. S. S. Sarma, R. D. Gulati, R. L. Wallace, S. Nandini, H. J. Dumont & R. Rico-Martínez Advances in Rotifer Research     

Comparison of colony foundation success between sexual pairs and female asexual units in the termite Reticulitermes speratus (Isoptera: Rhinotermitidae)

In termites, a male and a female usually found a colony cooperatively. However, pairing efficiency tends to be low in Reticulitermes speratus because of a limited mate-searching range, the female-biased sex ratio, and a relatively low calling ability. Females that fail to pair with males found colonies either in female–female pairs or even alone. In the laboratory, we examined colony foundation by single females (F), female–female pairs (FF), and normal male–female pairs (FM). The time until colony foundation (when termites began excavating wood baits) differed significantly among the unit types. Time until excavation was much longer for single females than for FF and FM units, which reflects the relative success of colony foundation. The survival rate of single females was also significantly lower than that of FF- and FM-unit females, although there was no difference between FF and FM units. This result demonstrates that cooperation, even female–female, promotes female survivorship. Nevertheless, the number of progeny per female was significantly lower in FF units than in FM units, possibly because females of FF units must share reproductive output. These results lead us to the conclusion that a normal monogamous pair is the best unit for colony foundation. Nevertheless, females alone can establish colonies by parthenogenesis, and even female–female cooperation promotes colony foundation success if pairing with males is not possible. Considering the functional decision for females in F and FF units of how much time to spend searching for a male mate, we believe that these facultative pathways of colony foundation by parthenogenesis have adaptive significance. Received: November 30, 2000 / Accepted: March 6, 2001     

Chemosensory cues allow male Tenebrio molitor beetles to assess the reproductive status of potential mates

Males of many insect species, including beetles, choose their mates according to their reproductive status. However, the ways in which male beetles evaluate female reproductive status have received little attention. We tested the existence of male mate choice in the mealworm beetle, Tenebrio molitor, by observing mating and courtship behaviour of males given simultaneous access to pairs of females differing in their reproductive status: (1) mature versus immature; (2) virgin versus previously mated; (3) familiar (mated with the experimental male) versus unfamiliar (mated with a different male). Males courted and mated preferentially with mature and virgin females. To determine whether chemical cues played a role in these discriminations, we exposed males to filter paper squares bearing chemical cues from different types of females: (1) virgin versus mated; (2) mature versus immature. Males were significantly more attracted to those squares bearing chemical cues from virgin and mature females, suggesting that males can assess female reproductive status on the basis of chemical cues alone.     

Mate choice and competition in the barklouseLepinotus patruelis (Psocoptera: Trogiidae): The effect of diet quality and sex ratio

Courtship varies among individuals, partly because individuals differ in quality. To explore proximate factors affecting courtship behavior, I investigated the effect of diet quality on mate choice and competition in the barklouseLepinotus patruelis Pearman (Psocoptera: Trogiidae) in the laboratory. The effect of sex ratio on mate choice was also addressed. Some males were found to exhibit active mate choice, and rejected females in both male- and female-biased sex ratio groups, although they were more likely to do so in a female-biased sex ratio group. Diet quality affected male mate choice: males on high-quality diets were significantly more likely to reject females than males on low-quality diets. Males exhibited choice significantly more often than females, who showed no overt signs of choosiness. Both males and females competed for, access to mates: both sexes attempted to interfere with mounted pairs and females grappled. The choosiness of the male may have directly affected the incidence of female competition. The results also suggest that the patterns of mate choice inL. patruelis differ from those expected by conventional sex role theory.     

Mating order-dependent female mate choice in the polygynandrous common lizard Lacerta vivipara

Recent studies indicate that directional female mate choice and order-dependent female mate choice importantly contribute to non-random mating patterns. In species where females prefer larger sized males, disentangling different hypotheses leading to non-random mating patterns is especially difficult, given that male size usually correlates with behaviours that may lead to non-random mating (e.g. size-dependent emergence from hibernation, male fighting ability). Here we investigate female mate choice and order-dependent female mate choice in the polygynandrous common lizard (Lacerta vivipara). By sequentially presenting males in random order to females, we exclude non-random mating patterns potentially arising due to intra-sexual selection (e.g. male–male competition), trait-dependent encounter probabilities, trait-dependent conspicuousness, or trait-dependent emergence from hibernation. To test for order-dependent female mate choice we investigate whether the previous mating history affects female choice. We show that body size and body condition of the male with which a female mated for the first time were bigger and better, respectively, than the average body size and body condition of the rejected males. There was a negative correlation between body sizes of first and second copulating males. This indicates that female mate choice is dependent on the previous mating history and it shows that the female’s choice criteria are non-static, i.e. non-directional. Our study therefore suggests that context-dependent female mate choice may not only arise due to genotype-environment interactions, but also due to other female mating strategies, i.e. order-dependent mate choice. Thus context-dependent female mate choice might be more frequent than previously thought.     

Promiscuous tendencies in the round-eared sengi: a male’s perspective

Although monogamy is interpreted as risk-adverse strategy by reducing intra-sexual conflicts, most pair-living males increase their reproductive success by engaging in extra-pair copulations. However, little is known about costs involved in such extra-pair attempts from the male’s perspective. We investigated promiscuous tendencies of paired Macroscelides proboscideus (round-eared elephant-shrew or round-eared sengi), a pair-living small mammal occurring in southern Africa. In particular, we measured potential costs of extra-pair attempts for the males. For this, we conducted laboratory experiments, involving interactions between neighbouring male–female pairs. Data collection included direct behavioural observations and establishing the morphological and physiological characteristics of males. Both sexes intruded into the neighbouring area, and initiated sexual behaviour with neighbours of the opposite sex. Males which displayed a higher marking frequency in the neighbouring area received more sexually motivated behaviour from neighbouring females. Resident males attacked intruding males. Aggression experienced by and marking behaviour of intruding males was positively correlated with body mass losses. Furthermore, glucocorticoid levels, determined from analyses of faeces and urine samples, correlated positively with body mass losses and marking behaviour of intruding males, indicating costs of intruding. We conclude that male as well as female round-eared sengis have promiscuous tendencies. Although interactions with neighbouring females may offer potential benefits for males in terms of enhanced reproductive success, males also seem to pay substantial costs, as indicated by body mass losses and elevated stress hormone levels.     

Opportunity for sexual selection arises at moderate densities in the frog Rana chensinensis: an experimental approach

Heavily male-biased operational sex ratio and high male density characterizing explosively breeding anurans are expected to enhance the opportunity for sexual selection through large-male mating advantage brought about by severe male–male competition. However, our previous study on the temperate frog Rana chensinensis suggested that these two factors might restrict the opportunity under natural conditions for females to be grasped by any male they meet first, and scramble competition between males occurred less frequently because amplectant pairs hid on the bottom of ponds. In the present experimental study, in which the operational sex ratios and male densities were much lower and individual interactions were sufficiently longer than in natural population, we found a large-male pairing advantage in female mate choice, multiple mate choice, and male displacement experiments. These findings lend circumstantial support for the predication, although there was no evidence of large-female mating advantage in a male mate choice experiment.     

A review of the potential of surface flow constructed wetlands to enhance macroinvertebrate diversity in agricultural landscapes with particular reference to Integrated Constructed Wetlands (ICWs)

Discriminating female mate preferences enhance the variance in reproductive success among males of a population and create a potential for sexual selection, which can account for trait evolution and diversification. Fish color patterns are among the prime targets of mate choice-driven sexual selection. Populations of the cichlid Tropheus from Lake Tanganyika display remarkable geographic color pattern variation, but the role of female choice in their rapid and rich phenotypic diversification is unclear. Males and females establish a pair bond prior to spawning monogamously, but as brood care is strictly maternal, female investment in reproduction is high and the operational sex ratio is male-biased. Therefore, variance in male reproductive success can accrue if individual males succeed repeatedly in securing a mate. To test this prediction in the red colored Tropheus moorii “Chimba”, four pairs of males were presented to a series of females and female mate preferences were inferred from pairwise interactions. There was a significant difference in mating success between the males of each pair (P < 0.001 over all trials), as—with one exception—females shared preferences for the same males. Male courtship activity was strongly correlated with female choice. Our experiment suggests that female choice contributes to the variance in male reproductive success in the tested population.     

Pair-clinging behavior by males ofAtrophaneura alcinous (Lepidoptera: Papilionidae)

Females ofAtrophaneura alcinous usually mate soon after eclosion. Their ostium bursae becomes plugged with male secretion which reduces chances of remating. Males frequently cling to a copulating pair and wait for completion of copulation. This was observed in 66% of 198 copulating pairs, with a maximum of 5 males clinging at one time during the course of a copulation. Males clinging for longer periods were more successful in copulation with the freshly mated female than those clinging for shorter periods. Despite the plugging effect, females may mate more than once. Clinging males were responsible for 61% of re-copulations and 53% of re-inseminations. Clinging behavior may be regarded as an effective male mating strategy to exploit freshly mated females, and an alternative to finding virgin females.     

Are females of Ilyoplax pusilla (Brachyura: Dotillidae) attracted to groups having more waving males?

Males of the dotillid crab Ilyoplax pusilla wave at approaching females during the breeding season. They also, however, often perform waving that is not directed toward any particular individual. This undirected waving is associated with the presence of male neighbors and may function in male–male competition. It may also, however, act as a long-range female attractant. To test whether undirected waving functions to attract females, we conducted a field experiment that manipulated the abundance of waving males. We found that females preferred to approach groups that had more waving males. This suggests that undirected waving by male I. pusilla functions as a long-range courtship signal.     

Female choice, male interference, and sperm precedence in the red-spotted newt

Darwin first identified female choice and male—male competitionas forms of sexual selection resulting in the evolution of conspicuoussexual dimorphism, but it has proven challenging to separatetheir effects. Their effects on sexual selection become evenmore complicated when sperm competition occurs because spermprecedence may be either a form of cryptic female choice ora form of male—male competition. We examined the effectsof tail height on male—male competition and female choiceusing the sexually dimorphic red-spotted newt (Notophthalmusviridescens viridescens). Experiment 1 examined whether maletail height influenced male mating success. Males with deeptails were more successful at mating with females than thosewith shallow tails. Successful, deep-tailed males also were bigger(snout-vent length; SVL) than unsuccessful, shallow-tailed males,but they did not vary in tail length or body condition. Of these,only tail height and tail length are sexually dimorphic traits.Experiment 2 tested the hypothesis that the differential successof males with deeper tails was due to female choice by examiningboth simultaneous female preference for association and sequentialfemale choice. We found no evidence of female choice. When maleswere not competing to mate with females, tail height did notinfluence male mating success. Successful males did not havedifferent SVL and tail lengths than unsuccessful males. Thus,tail height in male red-spotted newts appears to be an intrasexuallyselected secondary sexual characteristic. Experiment 3 usedpaternity exclusion analyses based on molecular genetic markersto examine the effect of sperm precedence on sperm competitionin doubly-mated females. Sperm precedence likely does not havea pervasive and consistent effect on fertilization success becausewe found evidence of first, last, and mixed sperm usage.     

Mutual mate choice by mountain pine beetles: size‐dependence but not size‐assortative mating

1. Mutual mate choice may be rare, occurring when both sexes invest heavily in reproduction, mating opportunities are abundant, and individuals differ in quality. 2. Mountain pine beetles, Dendroctonus ponderosae (Curculionidae: Scolytinae) appear to meet the conditions for mutual mate choice. We introduced males to females in breeding sites and observed the occurrence and speed of a male entering a female's gallery. We tested for consequences of mutual mate choice, namely condition‐dependent choosiness and assortative mating. 3. Males were more likely to enter a female's gallery when the gallery was in a smaller tree with less resin production and when the gallery was larger. Female body size and condition did not influence the probability of entry. Larger males were less likely to enter a gallery than were smaller males, probably because of size‐dependent choosiness rather than physical limitations. 4. Small males took longer to enter galleries of large females than of small females, whereas large males entered as quickly into galleries of large females as small females. This suggests size‐dependent choosiness by females. 5. No assortative mating by body size was detected, probably because males appeared to choose on the basis of female‐associated resources rather than on female traits.     

Courtship tactics by male Ilyoplax pusilla (Brachyura, Dotillidae)

Mating in the dotillid crab Ilyoplax pusilla occurs after the female enters the male’s burrow in the tidal flat. Males use two tactics to cause females to enter their burrows for mating: the male either directs claw waving to the female (courting-wave display), to which the females responds by following the male to his burrow, or the male runs rapidly away from, then back toward, his burrow (dash-out-back display), which startles the female into his burrow. Males more often used the courting-wave than the dash-out-back display, but mating success did not differ between the two tactics. Male use of either tactic was influenced by date, female density and male size; the courting-wave display was used by larger males, later in the breeding period, and under higher female density.     

Male pioneering as a mating strategy: the case of the beetle Maladera matrida

1. The phenomenon of male pioneering, whereby males are active (daily) before females, is compared with protandry, a phenomenon whereby males are sexually matured before females. The effects of natural selection and sexual selection on both phenomena are discussed. 2. In Maladera matrida Argaman (Coleoptera: Scarabaeidae), males emerged from the soil a few minutes before females, every evening, to feed and mate. 3. Aggregations of males and females are formed at sites of plant damage where males initiated feeding. 4. All copulations were initiated at the beginning of the activity period so males that emerge earlier may increase their probability of finding a mate. 5. Females mate only once in an evening. 6. Males that emerge later may not find receptive females with which to mate. 7. The aggregation of beetles per se did not contribute to the individual mating success because it occurred during and after the copulating had begun. 8. Male pioneering may evolve through sexual selection by means of male–male competition in time to achieve receptive females.     

Intrasexual Competition and Mating Behavior in <Emphasis Type="Italic">Ptomascopus morio</Emphasis> (Coleoptera: Silphidae Nicrophorinae)

In nicrophorine beetles, genus Nicrophorus care their larva using small vertebrate carrion, whereas genus Ptomaucopusreproduce with small vertebrate carrion but show no parental care. Aggression and sexual behavior were examined in Ptomascopus morio and Nicrophorus quadripunctatus. Nicrophorus quadripunctatus had intense female–female as well as male–male contests. In Ptomascopus morio, by contrast, female–female aggression was rarely observed. Male–male aggression (pushing, biting, male–male mounting) in Ptomascopus morio was observed when a resource for breeding was present, whether or not a female was present. The lack of female–female aggression, and male–male aggression when resources but not females are present, suggest that the mating system of Ptomascopus morio is resource defense polygyny. Large males of Ptomascopus morio were also found to exhibit mate choice, preferring large females over small females.     

Group unity of chimpanzees elucidated by comparison of sex differences in short-range interactions in Mahale Mountains National Park,Tanzania

Chimpanzees (Pan troglodytes) form multi-male and multi-female unit groups with fission–fusion grouping patterns. Short-range interaction (SRI) plays an important role in the unity of these groups and in maintaining social bonds among members. This study evaluated three models of chimpanzee social structure that differed according to the emphasis each placed on social bonds between the sexes, i.e., the male-only, the bisexual, and the male-bonded unit-group model. I investigated differences in SRI between the sexes among group members in well-habituated wild chimpanzees in Mahale Mountains National Park, Tanzania. I followed six focal adult males and six females, and quantified their respective SRI with other chimpanzees. Except between subordinate males and adult females, adults in general engaged in SRI with about 60–90% of the individuals with whom they made visual contact each day, whether in large or small parties. Although the number of social grooming (SGR) partners was limited, male–male SGR networks were wider than were either male–female or female–female SGR networks among adults. The number of contact-seeking behavior (CSB) partners was also limited, but dominant males had more CSB partners. Adult females mainly interacted by pant-grunt greeting (PGG) with adult males, but tended to do so mainly with the highest-ranking male(s) within visual contact. These results indicated that the social bonds among adult males were essential to group unity. Because of clear male dominance, adult females established peaceful coexistence with all group members despite less frequent SRI with subordinate males by maintaining affiliative social bonds with dominant males, thereby supporting the male-bonded unit-group model. Adult females had many female SRI partners, but these interactions did not involve performing conspicuous behaviors, suggesting that females maintain social bonds with other females in ways that differ from how such bonds are maintained with and between adult males.