Multiple signals in the palmate newt: ornaments help when courting

There is increasing evidence that female mate choice is often based on the assessment of multiple male traits, involving both morphology and behavior. We investigated female mate choice for multiple male traits in the palmate newt, Lissotriton helveticus, including male tail filament length, hind foot web size, crest development, body size, ventral coloration, and courtship display activity. Observations of courtship display in the field revealed that females spent more time in front of males with longer tail filaments. Laboratory experiments revealed a more detailed relationship between filament length and courtship display. We found that females took more sperm masses from males with both longer filaments and greater display activity. Experimental shortening of the tail filament length substantially decreased the number of male sperm masses transferred. However, when we experimentally reversed relative filament length between two males in mating trials, male mating success was explained by courtship activity and not by filament length. Our results show that female palmate newts value multiple traits during mate choice, including both morphological ornaments and reproductive behaviors in males. Our results further suggest that, when filament length is below a certain threshold, females may value the information content of courtship activity over that of filament length.     

Effect of temperature,salinity and food level on sexual and asexual reproduction in Brachionus plicatilis (Rotifera)

The reproductive response of sexual and asexual female Brachionus plicatilis (Muller) was examined over temperatures ranging from 20° to 40°C, salinities from 5 to 40 S, and food levels from 0.25 to 20 g Chlorella vulgaris dry-weight per ml. Reduced food levels, as well as temperature and salinity extremes, reduced reproduction of both sexual and asexual females, but did so differentially. Reproduction by sexual females was reduced to a greater extent at environmental extremes than asexual females. The broad, flat reproductive response curve of asexual females extended beyond the limits of the narrower, more sharply peaked curve of sexual females. Thus zones of exclusively asexual reproduction exist at environmental extremes where sexual reproduction is physiologically restricted. These results are corroborated by a comparison of the lifetime fecundity of individual sexual and asexual females over a 20°C temperature range. No differences in lifetime fecundity occurred between sexual and asexual females at 18° and 28°C. At 38°C, however, asexual female fecundity reached its highest level, while sexual female fecundity declined 15%. The appearance of sexual females in rotifer populations in the result of both inducible and repressible factors.     

A reliable indicator of female fecundity: the case of the yellow belly in<Emphasis Type="Italic"> Knipowitschia panizzae</Emphasis> (Teleostei: Gobiidae)

A large body of research has developed on male ornaments and their role in sexual selection while female ornaments have been less investigated. This study focused on the presence, function, and possible role of a sexually dimorphic nuptial trait as an indicator of female quality in a fish species presenting conventional sex roles: the yellow patch on the belly of Knipowitschia panizzae females. Aquarium spawning experiments demonstrated that this ornament is due to dermal pigments, is displayed only when the female is ready to spawn, and is switched off within minutes from the end of egg deposition. This sexual trait is variable in size among females and indicates female fecundity relative to her own body size. As a consequence, female yellow belly appears to be a reliable indicator of female quality as measured by current fecundity. Field data on natural nests highlights that males usually perform parental care on only one egg batch at a time. The modality of egg deposition suggests that males may be limited in their potential reproductive rates by environmental factors. Male limitations in egg care could favour male choosiness and the evolution of female nuptial displays.Communicated by R. Cattaneo-Vietti, Genova     

Influence of mating order on courtship displays and stored sperm utilization in Hermann's tortoises (Testudo hermanni hermanni)

Sperm storage is widespread in many vertebrate groups, and it is frequently associated with promiscuous mating systems. Chelonian species are one of the most outstanding examples of a promiscuous group capable of long-term sperm storage; specialized structures have evolved within the oviducts of these vertebrates to ensure sperm vitality across reproductive cycles. Thus far, few studies have investigated the factors regulating multiple paternity, sperm usage by females and paternity distribution in successive clutches. This study aimed to investigate the effect of mating order on male mounting and reproductive success in Testudo hermanni hermanni, combining behavioural and genetic data. A series of planned matings were performed, within which experimental females were mated sequentially to two different males under controlled conditions. Observations conducted during experimental matings revealed that courtship displays did not significantly differ between the first and second males to mate with a female. Interactions ending with a mount were characterized by a significantly higher intensity and occurrence of determinate courtship behaviours, for example biting and running after the female. Paternity analysis performed on hatchlings produced from experimental females revealed that 46 % of the clutches were multi-sired. A significant contribution of previous years' partners was still found, thus confirming the long-term sperm storage within the female oviduct in this species. Finally, mating order did not significantly affect the reproductive success of experimental males during the on-going reproductive season.     

Leg ornamentation and the efficacy of courtship display in four species of wolf spider (Araneae: Lycosidae)

This study used both correlative and experimental video playback methods to test the hypothesis that the secondary sexual traits of male wolf spiders act to increase the efficacy of visual courtship displays. Direct observations of courtship of several lycosid genera and a review of the literature revealed a significant association between ornamentation and visual courtship displays. This suggests that the ornamentation may be playing the role of amplifier for a visual display. To test this hypothesis, male courtship behaviors of four Schizocosa species were experimentally manipulated using video-imaging techniques. Females of species with non-visually displaying, non-ornamented males (Schizocosa duplex and S. uetzi) did not increase in frequency of receptivity when tufts were added to conspecific males. In a species with a visual display and foreleg pigmentation (S. stridulans), the addition of foreleg tufts increased female receptivity. In a tufted species (S. crassipes), females tended to decrease their receptivity when male ornamentation was completely removed. In visually displaying species, ornamentation acts to increase female receptivity, supporting its role as an amplifier of a visual display. Received: 29 December 1997 / Received in revised form: 23 October 1999 / Accepted: 13 December 1999     

Reproductive cycle of<Emphasis Type="Italic"> Bispira volutacornis</Emphasis> (Polychaeta: Sabellidae) on the west coast of Ireland

The reproductive cycle of Bispira volutacornis, a dioecious polychaete, was followed over two spawning periods in Lettercallow Bay, on the west coast of Ireland. In the younger generation of sexually mature adults, gender can be determined on the basis of colour, the dorsal aspect of the males being cream at maturation, whilst the females are pink at the previtellogenic stage and an olive green at maturation. An annual reproductive cycle of 12 months shows that oocyte/sperm maturation is extraovarian. The male reproductive cycle proceeded at a more rapid pace than that of the female cycle, with sperm maturing earlier. Spawning in B. volutacornis occurred between late July and early September. B. volutacornis is a polytelic species, surviving spawning and releasing all gametes in a single batch, with a mean spawned oocyte diameter of 118 m. The reproductive cycle appears to be regulated by a seasonal change in water temperature, and spawning is timed to produce larvae during the autumnal planktonic bloom in Lettercallow Bay.Communicated by J.P. Thorpe, Port Erin     

Mate choice in the yellow-shouldered widowbird: correlates of male attractiveness

Recent investigations of male ornaments in sexual selection have used experimental manipulation of tail length in three widowbird species, but only for one of these have correlates of male reproductive success been reported. I examined correlates of male attractiveness to nesting females over two breeding seasons for the polygynous yellow-shouldered widowbird, Euplectes macrourus, in order to discover which cues females may be using to select mates. The black, long-tailed ( 10 cm) males defend large territories and build nest frames, or cock's nests, which females then line and use for nesting. I examined various aspects of male morphology, five behavioral displays, territory characteristics, and the number of cock's nests that males built. Few correlates of mating success were found. The best predictor was the number of cock's nests that a male builds, though one courtship display also correlated with male mating success in 1 year, as did average grass height. Tail length did not correlate with male mating success. A partial correlation analysis confirmed that cock's nests and, in 1 year, grass height, were the primary contributers to male success. Females may choose where to nest primarily on the availability of suitable nesting sites. Long tails may be used by females seeking extra-pair copulations or in male-male competition for territories.     

Sexual selection,male morphology,and the efficacy of courtship signalling in two wolf spiders (Araneae: Lycosidae)

Males of the brush-legged wolf spider, Schizocosa ocreata (Araneae: Lycosidae), possess a conspicuous male secondary sexual character: dark pigmentation and tufts of bristles on the tibiae of their forelegs. We tested several hypotheses relating to the role of this conspicuous trait in sexual selection. Triad mating experiments suggest that the tufts do not play an obvious role in the operation of sexual selection by either male competition or female choice, as there were no significant differences in the mating success of intact and experimentally shaved males. However, females mated more often with males that initiated courtship first, suggesting that capture of a female’s attention by male signalling may play a critical role. In behavioral experiments that paired a single male with a female in arenas that allowed both visual and vibratory signal transmission during courtship, female receptivity did not vary significantly with the presence or absence of tufts. However, experiments that isolated the visual component of communication (by eliminating vibratory communication) revealed a significant effect of the presence of tufts: females showed receptivity less often to males with tufts removed. Female response to visual signals was much greater in S. ocreata than in its sibling congener, Schizocosa rovneri, which lacks male tufts. We hypothesize that the tufts serve to increase the efficacy of visual displays of S. ocreata, as vibratory communication is constrained by the complex leaf litter habitat of some populations. Such environmental constraints may make visual signalling over distance a critical factor for effective courtship communication, which may in turn strongly influence male fitness. Received: 30 September 1994/Accepted after revision: 4 August 1995     

Dart receipt promotes sperm storage in the garden snail Helix aspersa

During courtship, many helicid snails attempt to pierce the body walls of their mating partners with mucus-coated calcareous darts. The mucus covering the dart induces conformational changes in the female reproductive tract of the recipient, closing off the entrance to the gametolytic bursa copulatrix. We have tested the effect of dart receipt on the number of sperm stored by once-mated snails, Helix aspersa. Snails that were hit by darts stored significantly more sperm than did snails that were missed. Additionally, the effect of the dart was stronger in smaller animals and the number of sperm stored decreased with the shell volume of the recipient. Although larger animals produced larger spermatophores, dart-shooting success was not related to the number of sperm transferred. These data suggest a role for dart-shooting in post-copulatory sexual selection.     

Male remating and female fitness in the wolf spider Pardosa astrigera: the role of male mating history

Although the effects of male mating history on female reproductive output and longevity have been studied in insects, few such studies have been carried out in spiders. In a mating system in which females are monandrous while males are polygynous, females may incur the risk by mating with successful males that have experienced consecutive matings and suffer from the possible depletion of sperm and/or associated ejaculates. Here, we examine the effects of male mating history on male courtship and copulation duration, female reproductive fitness, and female adult longevity of the wolf spider, Pardosa astrigera. Results indicated that male mating frequency had little effect on their subsequent copulation success, and of 35 males tested, about half of the males were able to copulate with five virgin females successively at an interval of 24 h. Male mating history had little effect on their courtship duration. However, male mating history significantly affected male copulation duration, female adult longevity, and reproductive output. Males that mated more frequently copulated longer and more likely failed to cause their mates to produce a clutch, although there was no significant difference in the number of eggs laid and the number of eggs hatched regardless of the first clutch or the second one. Multiple mating of male P. astrigera resulted in significant reduction in female adult longevity. Our results indicate that monandrous females mating with multiple-mated males may incur substantial fitness costs.     

Female courtship in the Banggai cardinalfish: honest signals of egg maturity and reproductive output?

Despite the vast literature on male courtship behaviour, little is known about the function and information content of female courtship behaviour. Female courtship behaviour may be important in many species, particularly where both sexes invest heavily in the offspring, and if such behaviours contain honest information regarding a females potential reproductive investment, they may be particularly important in male mate choice. Using observations of two female courtship behaviours (the rush and the twitch) from experimental pairings in the Banggai cardinalfish (Pterapogon kauderni), I addressed the question of whether these courtship behaviours contained information on female reproductive output (clutch weight) and egg maturity (proximity to spawning), traits commonly associated with male mate choice. I especially focused on the importance of these courtship behaviours in relation to other female characters, such as size and condition, using multiple regression. I found that one of these behaviours, the rush, was strongly associated with fecundity, whereas size, condition and the twitch were not. Further, I found that the twitch behaviour was associated with how close to actual spawning a female was. The results suggest that female courtship behaviour may convey highly important information in a mate choice context. I discuss the adaptive value of honest information in female courtship behaviour in light of these results.Communicated by K. Lindström     

Do microsporidian parasites affect courtship in two-spotted gobies?

Courtship and body condition of male two-spotted gobies (Gobiusculus flavescens) harbouring naturally acquired microsporidian infections were compared with those of microsporidian-free males under standardized conditions in the lab. Although parasite infection had no apparent effect on individual condition, it significantly affected male courtship intensity. This effect, however, was affected by the intensity of the female courtship. Our study is one of the first to demonstrate a sub-lethal behavioural effect of a microsporidian parasite that could negatively affect the reproductive success of infected individuals. Our results also suggest that secondary sexual traits like courtship may be more sensitive to moderately detrimental effects of parasite infection than classical condition indices.     

Experience with chemotactile cues indicating female feeding history impacts male courtship investment in the wolf spider <Emphasis Type="BoldItalic">Schizocosa ocreata</Emphasis>

Male fitness is often determined by the ability of the male to gain access to multiple mates, although in species that exhibit sexual cannibalism, males might increase their likelihood of being cannibalized with each encounter. This risk should create selection for males who are able to perceive potential risks associated with mating encounters. We studied male Schizocosa ocreata wolf spiders to determine whether they use female chemotactile cues (silk, excreta) as indicators of potential risks and how these cues affected subsequent male courtship behaviors. Female treatments included satiated females vs. starved females, as well as a treatment where females had recently cannibalized a male S. ocreata. We performed experiments to assess (1) if males use female chemotactile cues to determine potential risks associated with differing female feeding treatments and alter courtship investment, and 2) how male experience with female chemotactile cues affected courtship investment in subsequent female encounters. At first encounter, males do not vary courtship investment (number of bouts, duration, and vigor) with different female feeding treatments. However, male behaviors during subsequent encounters with female chemotactile cues varied in complex ways, depending on female feeding treatment and male first encounter experience. These data suggest that male experience impacts perception of female chemical cues and offers the male opportunity to avoid sexual cannibalism in high-risk situations.     

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.     

Rhythmic male reproductive behavior controls timing of courtship and mating in Laupala cerasina

In many organisms, mating behavior occurs at a particular time of day, which may be important for avoiding mate competition or interspecific mating. Crickets of the Hawaiian genus Laupala exhibit an unusually protracted courtship in which males produce a series of nuptial gifts prior to the species-typical time of mating. Mating time is one of several rhythmic behaviors that have diverged among closely related Laupala species, which exhibit an extremely high speciation rate. Mating rhythm may reflect direct selection on male and/or female sexual receptivity or the pleiotropic consequence of selection on other rhythmic behaviors. To examine the role of sexual rhythmicity in Laupala cerasina, we characterized the time boundaries or “circadian gate” of courtship and mating, as well as female phonotactic response to male song. We also examined which sex is responsible for mating rhythmicity by phase-shifting males relative to the female photophase. Our results demonstrate that mating behavior is gated by the end of the light phase. Time limits to female mating receptivity were not observed and thus male rhythm alone appears to be responsible for the timing of mating. Furthermore, when courtship is initiated later in the day, males produce fewer nuptial gifts and increase nuptial gift production rate while delaying mating, suggesting that the number of gifts a female receives is important to male reproductive success.     

Correlates of male reproductive success in the browncheek blenny,Acanthemblemaria crockeri (Blennioidea: Chaenopsidae)

Summary Acanthemblemaria crockeri exhibits a resource defense polygyny mating system with male parental care. Females preferred to mate with the larger of two males in laboratory experiments, and male size was positively correlated with the number of eggs defended by males at two sites in the Gulf of California, Baja California, Mexico. Females appeared to avoid mating with males defending heavily-fouled shelters. The role of other factors including the intensity of male courtship coloration and displays in determining male reproductive success was studied in Bahia San Carlos, Sonora, Mexico by providing glass vials as shelters. This provided a non-destructive technique for counting of eggs defended by males and allowed repeated assessment of the reproductive success of individual males. Males varied greatly in their duration of residency of shelters and in their color score and intensity of courtship displays. The number of eggs received by males was positively correlated with their duration of residency in shelters, and females appeared to avoid mating with new residents. Mated males deserted shelters less frequently and were more likely to receive future matings than were unmated males. The mean color score of males was unrelated to their reproductive success, while the intensity of male courtship displays was negatively correlated with the number of eggs received. This may have resulted because female mate choice is based on multiple criteria, including some that more accurately reflect the quality of parental care afforded by males.     

Control and coordination of reproduction and molting in the spiny lobster Panulirus argus

Behavioral and physiological correlates of vernal reproduction and molting were investigated experimentally in laboratory-held spiny lobsters (Panulirus argus Latreille) from Florida to determine their control by photoperiod and temperature. Behavioral measures included courtship, copulation and aggression; physiological measures included the timing and frequency of ecdysis, spermatophore emplacement, oviposition, egg hatching, and setal (pleopod) and gonadal development. Behavior resembled that observed in nature, as evidenced in the expression of complete reproductive cycles. Long daylengths and warmer temperatures enhanced courtship, spawning frequencies, and female gonadal development, although gonadal recrudescence occurred in large females irrespective of photoperiod. A significant interaction effect between photoperiod and temperature indicated a need to examine joint, non-independent factor effects upon reproductive parameters. Photoperiod and temperature did not significantly influence aggression and male gonadal development. Photoperiod did not affect molting rates significantly. However, warm temperatures enhanced molting and growth such that sub-adult and young adult lobsters (both sexes) had highest molting rates, followed by large adult males, which in turn had higher rates than reproductively active females. Female setal development correlated positively with female size, and changed abruptly at ecdysis. Although setal lengths were highly variable, setae of 9 mm or more signified sexual maturity. Reproduction and molting were size-dependent: the largest adults mated or spawned, the smallest molted, and intermediate-sized ones either mated, or molted before mating. Spiny-lobster reproductive and molting patterns were influenced by photoperiod and temperature in a complex manner depending upon sex, season, size and developmental state.     

Sexual selection and cues for female choice in leks of Jackson's widowbird Euplectes jacksoni

Summary Sexual selection through female mate choice was investigated in the lekking Jackson's widowbird by applying multivariate selection analysis to observational data from four leks. Males perform a stereotyped jump display on small display courts (dance rings) constructed by the males in open grassland. Females visit the lek solely for mating and nest on their own, away from the lek area. Few cases of interference during courtship and absence of position effects on mating success indicated that female choice within the leks was not pre-empted by male-male competition. In a set of 11 male traits with mating success as the dependent fitness measure, significant selection differentials (covariances) were found for the length of the conspicuous tail and the rate of the jump display, suggesting sexual selection of these traits. They also showed the largest selection gradients (partial effects) and thereby seem to be the cues on which females base their choice. The success of males in obtaining copulations appears to depend on two components: display rate and lek attendance affect the number of female visits, whereas tail length seems to primarily influence the chance of copulating with a visiting female. Tail length was positively related to a measure of body condition, which is of interest with regard to the suggestions that sexual ornaments may serve as indicators of male viability.     

Effects of male dominance and courtship display on female choice in the ring-necked pheasant

Male traits and behaviours acting in mate choice and intrasexual competition are expected to be congruent. When studying their evolution, this often makes it difficult to differentiate between these two components of sexual selection. Studies are therefore needed on mate choice in conjunction with the role of displays and dominance. We present the results from two experiments conducted to investigate the effects of male dominance and courtship displays on female choice in the ring-necked pheasant, Phasianus colchicus, controlling for differences in morphological male traits. We found: (1) different courtship behaviours had different effects on female choice: females were mainly attracted by the feeding courtship behaviour, while another courtship display (the lateral display) was effective in producing the copulation-acceptance response by the females; (2) subordinate males performed the courtship behaviour before females less frequently than dominant males, and females reinforced intrasexual selection by choosing dominant males, and (3) subordinate males in visual contact with a dominant became less attractive to females. The results support the idea (armament-ornament model) that female pheasants may benefit from using traits selected in male-male competition as clues for mate choice. Received: 23 October 1997 / Accepted after revision: 7 October 1998     

Why do male Callosobruchus maculatus beetles inseminate so many sperm?

Male Callosobruchus maculatus F. (Coleoptera: Bruchidae) inseminate more sperm than females can effectively store in their spermathecae. This study examines the adaptive significance of excess sperm transfer by measuring components of male and female reproductive success in response to manipulating the number of sperm inseminated. The number of sperm transferred during copulation was reduced from 56,000 ±4,462 to 8,700±1,194 by sequentially mating males to virgin females. Reducing the number of sperm inseminated by the first male to mate had no effect on the extent of sperm precedence, but reducing the number of sperm inseminated by the second male resulted in a significant reduction in the extent of sperm precedence. When large numbers of sperm are inseminated the remating refractory period of females is increased. These results indicate that males transferring large numbers of sperm during copulation have a two-fold advantage at fertilization; they are more effective at preempting previously stored sperm and they are likely to father more offspring by delaying the time of female remating. The transfer of excess sperm does not appear to serve as nonpromiscuous male mating effort; the number of eggs laid, their fertility and the subsequent survival of zygotes were unaffected by manipulating the number of sperm inseminated. The underlying mechanisms of sperm precedence were also examined. Simple models of sperm displacement failed to accurately predict the patterns of sperm precedence observed in this species. However, the results do not provide conclusive evidence against the models but rather serve to highlight our limited understanding of the movement of sperm within the female's reproductive tract.