Quantifying male attractiveness and mating behaviour through phenotypic size manipulation in the Trinidadian guppy, Poecilia reticulata

Although many studies have examined the effects of male size on attractiveness and mating behaviour, few have taken genetic background into consideration. Phenotypic manipulation permits the experimental adjustment of morphological traits while keeping genetic background constant. Here, male guppies, Poecilia reticulata, an ideal model for this type of manipulation, were raised at different temperatures to produce sibling pairs that differed in size. These were then used to investigate male mating behaviour and male attractiveness, assessed through female mate choice, in relation to this size dimorphism. Further, male–male competition, which is intrinsic to male mating behaviour, is also likely to be affected by their size. Through the use of repeated measures analyses we demonstrate that females significantly prefer larger males and male size and competition significantly affect several aspects of male mating behaviour. Larger siblings perform more sneaky mating attempts and spend more time chasing females. The frequencies of both these behaviours increase with competition. While display frequency is unaffected by male size and competition, display duration and the amount of time spent attending females are reduced in the presence of competitors. This study highlights the use of phenotypic manipulation as a valuable tool for investigating behavioural interactions and confirms that both male size and competition are significant factors in the guppy mating system.     

Paternity and condition affect cannibalistic behavior in nest-tending bluegill sunfish

Theory of parental care evolution predicts that a parent should invest more in a brood when its fitness value is greater than alternative investments such as the parent's own survivorship or future broods. In fish, filial cannibalism (eating one's own offspring) is widespread and represents a challenge to parental care evolution. In this study, I investigated filial cannibalism in bluegill sunfish (Lepomis macrochirus). Bluegill are characterized by alternative mating tactics referred to as "parentals" and "cuckolders". Parentals delay maturation, construct nests, court females and provide sole parental care for the developing offspring. Cuckolders mature precociously and parasitize parentals using two tactics called "sneakers" and "satellites". I found that parentals that obtained fewer eggs during spawning appeared more likely to completely cannibalize their brood (total filial cannibalism: P=0.07), regardless of their condition. Among parentals that provided care, partial cannibalism was greater during the egg phase as compared to the fry phase of care, but it was unrelated to brood size. Throughout the care period, parentals in better condition were less likely to partially cannibalize their brood, indicating that parentals use cannibalism to replenish energy reserves. Independent of condition, parentals that were cuckolded more were more likely to eat part of their brood. This relationship was evident only after the eggs had hatched, which is consistent with data showing that parentals can use olfactory cues produced by fry but not eggs to assess their paternity. This latter result proposes that parentals may be selectively culling cuckolder offspring from their nest. These data provide empirical support for parental care theory, and the first evidence for the importance of paternity on cannibalistic behavior.Communicated by M. Abrahams     

Males in the shade: habitat use and sexual segregation in the Galápagos sea lion (Zalophus californianus wollebaeki)

In many sexually dimorphic species adult sexes tend to segregate socially, spatially, or in habitat use. Several hypotheses have been formulated regarding underlying mechanisms. We investigated terrestrial habitat use and sexual segregation in a tropical otariid, the Galápagos sea lion (Zalophus californianus wollebaeki), where most of the hypotheses can be ruled out a priori. Factors relating to thermoregulation and costs of locomotion were of prime importance for habitat use. Habitats directly adjacent to the sea, with simple structured flat surfaces, shade, and tide pools were most frequented, but sexes and age classes differed in their usage patterns. Sexual segregation, both spatial and by habitat was pronounced in the reproductive period (RP), but remained high during the nonreproductive period (NRP). A GLM model of habitat use showed that in both seasons adult males frequented habitat types that adult females and other age classes used much less. Males were most abundant in suboptimal inland habitats, which offered only shade for cooling. Females with newborns differed in habitat use from females with older offspring and lone females. Spatial and habitat segregation are explained most parsimoniously as by-products of social processes, primarily intrasexual competition and female avoidance of male harassment, linked to the polygynous mating system.Electronic Supplementary Material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Relatedness, polyandry and extra-group paternity in the cooperatively-breeding white-browed scrubwren (Sericornis frontalis )

We used DNA fingerprinting to examine the genetic parentage and mating system of the cooperatively breeding white-browed scrubwren, Sericornis frontalis, in Canberra, Australia. Our analyses revealed a remarkable variety of mating tactics and social organization. Scrubwrens bred in pairs or multi-male groups that consisted of a female and two or more males. Females were always unrelated to the pair male or alpha (dominant) male. Among multi-male groups we found three different mating tactics. Firstly, when alpha and beta (subordinate) males were unrelated, they usually shared paternity in the brood. This resulted in both males gaining reproductive benefits directly. Secondly, when beta males were not related to the female but were related to the alpha males, beta males sired offspring in some broods. In this situation, beta males gained reproductive benefits both directly and potentially indirectly (through the related alpha male). Thirdly, when beta males were related to the female or both the female and alpha male, they remained on their natal territory and did not sire any offspring. Thus beta males gained only indirect reproductive benefits. Overall, when group members were related closely, the dominant male monopolized reproductive success, whereas when the members were not related closely the two males shared paternity equally. This positive association between monopolization of reproduction and relatedness is predicted by models of reproductive skew, but has not been reported previously within a single population of birds. Other cooperatively breeding birds with both closely related and unrelated helpers may show a similar variety of mating tactics. Finally, we found that extra-group paternity was more common in pairs (24% of young) than in multi-male groups (6%), and we discuss three possible reasons for this difference. Received: 21 May 1996 / Accepted after revision: 14 December 1996     

Alternative mating behaviors of the queen polymorphic ant <Emphasis Type="Italic">Temnothorax longispinosus</Emphasis>

Mating behaviors of ants fall into two categories: female calling, in which a female alate releases pheromones that attract males, and male swarming, in which large male aggregations attract females. Female calling is common in species with queens that return to their natal nest to found colonies dependently after mating, while male swarming is common in species with queens that disperse to found independently. In some species that display both founding strategies, a queen-size polymorphism has evolved in which dependent-founding queens are smaller than independent-founding queens. Dependent founding is likely difficult if gynes (virgin queens) are mating in distant swarms. Therefore, a queen may adopt one or the other mating strategy based on its size and founding behavior. We investigated mating behaviors in the queen-polymorphic ant, Temnothorax longispinosus. Observations in laboratory mating arenas indicated that small gynes exhibited significantly lower flight activity than large gynes. Both forms mated in male swarms, and neither form exhibited female calling. The reduced flight activity of the small morph may facilitate returning to the natal nest after mating, provided the mating swarm is located nearby. Therefore, alternative colony-founding behaviors may be possible without the evolution of female-calling behavior; however, the reduced flight activity of small morphs may require that mating swarms are not distant from the natal nest.     

Dynamics of sperm transfer in the ant Leptothorax gredleri

Mating tactics differ remarkably between and within species of social Hymenoptera (bees, wasps, ants) concerning, e.g., mating frequencies, sperm competition, and the degree of male sperm limitation. Although social Hymenoptera might, therefore, potentially be ideal model systems for testing sexual selection theory, the dynamics of mating and sperm transfer have rarely been studied in species other than social bees, and basic information needed to draw conclusions about possible sperm competition and female choice is lacking. We investigated sperm transfer in the ant Leptothorax gredleri, a species in which female sexuals attract males by “female calling.” The analysis of 38 female sexuals fixed immediately or up to 7 days after copulation with a single male each revealed that the sperm is transferred into the female bursa copulatrix embedded in a gelatinous mass, presumably a spermatophore. Sperm cells rapidly start to migrate from the tip of the spermatophore towards the spermatheca, but transfer is drastically slowed down by an extreme constriction of the spermathecal duct, through which sperm cells have to pass virtually one by one. This results in the spermatheca being filled only between one and several hours after mating. During this time, the posterior part of the spermatophore seals the junction between bursa copulatrix and spermathecal duct and prevents sperm loss. The prolonged duration of sperm transfer might allow female sexuals to chose between ejaculates and explain previously reported patterns of single paternity of the offspring of multiply mated queens. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Reproductive trade-offs from mating with a successful male: the case of the tephritid fly Anastrepha obliqua

In lekking species, females may become sperm-limited when mating with sexually successful males, and this may be exacerbated by a poor male diet. Polygynous males may also be limited by the amount of accessory gland products (AGPs) they can transmit to females, which in turn may influence the females’ refractory period and longevity. Here, we tested the effect of male mating history, larval and adult diet on copula duration, mating intervals, female fecundity, fertilisation success, life span and likelihood to remate using sexually successful males of the lekking tephritid fly Anastrepha obliqua. Flies originated from either a native or exotic host fruit and were protein-fed or deprived. Male diet and larval host influenced copula duration, while the time elapsed between matings was affected by the interaction of mating order and male adult diet. Female fecundity was not influenced by female position in mating order or protein inclusion into the male diet. However, mating order and male larval diet influenced female fertilisation success. Importantly, as males mated successively they were less able to induce a refractory period on females, as the last females to mate with a male were more likely to remate and had slightly longer life spans than the first females to mate with males. These results might be attributed to a decrease in male AGPs with increasing male mating frequency. We discuss the role of conditional expression of male mating frequency with respect to A. obliqua’s life history, the trade-off that females face when mating with a successful male, the effect of larval diet on adult sexual performance and the possibility for sexual conflict to occur due to high male mating rates and fitness costs to females.     

Lifetime mating success, sexual selection and life history of fallow bucks (Dama dama)

We used data from a long-term study (15 years) of fallow deer to report for the first time the lifetime mating success, overall variance in lifetime mating success, and age-specific mortality levels of males. Fallow bucks that gain matings have higher social dominance rank, higher rates of fighting, and invest more in vocal display during the breeding season than unsuccessful males. Therefore, we examined if mating was associated with trade-offs in terms of survival, lifespan, and mating potential. We found that the variance in lifetime mating success was very high: 34 (10.7%) males mated, and of those, the 10 most successful males gained 73% of all matings (n=934). Mortality rates were generally high and only 22.3% (71/318) of males reached social maturity, i.e., 4 years. The oldest male was 13 years old. We found that fallow bucks that mated were not more likely to die during the following year, did not suffer from a reduction in lifespan, and did not incur lower mating success later in life as a result of mating during the early years of social maturity. Our results show that mating males at age 5 years (and possibly 9 years) may be more likely to survive than non-mating males. Additionally, the number of matings gained by males during the first years of social maturity was positively correlated with lifespan. We suggest that mating males are of higher quality than non-mating males because they are not more likely to incur trade-offs as a result of their increased reproductive efforts. Received: 9 November 1999 / Revised: 30 April 2000 / Accepted: 27 May 2000     

Sperm precedence, mating interval, and a novel mechanism of paternity bias in a beetle (Tenebrio molitor L.)

When females mate with more than one male, the ensuing sperm competition leads to the evolution of male mechanisms that skew paternity. Males of the yellow mealworm beetle (Tenebrio molitor) transfer a spermatophore to females during copulation, but sperm release and storage occur later. We investigated how the interval between two matings with different males affects sperm precedence by varying the interval between the copulations so that the second mating was either: (1) before sperm release from the first spermatophore (<5 min); (2) after sperm release but before spermatophore ejection (15–20 min); (3) after spermatophore ejection but before sperm storage (4 h), or (4) after complete sperm storage (24 h). We collected offspring over a period of 2 weeks and determined paternity by protein electrophoresis. There was second-male sperm precedence in all treatments, but when the interval was <5 min, the second male usually (86% of cases) had complete sperm precedence (i.e., P ₂=1). Investigations into the mechanism of second-male sperm precedence during <5-min mating intervals indicate that sperm release from the first spermatophore is inhibited, a phenomenon which has not been previously documented. Received: 31 January 2000 / Revised: 9 June 2000 / Accepted: 26 August 2000     

Sexual conflict in the snake den

Red-sided garter snakes (Thamnophis sirtalis parietalis) court and mate in spring, soon after they emerge from large communal overwintering dens in south-central Manitoba. Because of a massive bias in the operational sex ratio, every female attracts intense courtship from dozens to hundreds of males. We suggest that this courtship constitutes significant ”harassment,” because it delays the females’ dispersal from the den and hence increases their vulnerability to predation. Small females may face the greatest costs, because they are less able to escape from amorous males (who court all females, even juvenile animals). Our measurements show that males are stronger and faster than females. Experimental trials confirm that the locomotor ability of females (especially small females) is greatly reduced by the weight of a courting male. Arena trials show that intense courtship stimulates females to attempt to escape. Remarkably, some females that are too small to produce offspring may nonetheless copulate. This precocious sexual receptivity may benefit juvenile females because copulation renders them unattractive to males, and thus allows them to escape more easily from the den. Female ”tactics” to escape male harassment may explain other puzzling aspects of garter snake biology including size-assortative mating, temporal patterns in dispersal from the den, avoidance of communal dens by young-of-the-year snakes, and female mimicry. Hence, sexual conflict may have influenced important features of the mating system and behavioral ecology of these animals. Received: 8 May 2000 / Revised: 28 July 2000 / Accepted: 30 July 2000