Leigh W. Simmons

Analysing sperm competition data: simple models for predicting mechanisms

SummaryProspective models are developed for analysing sperm competition data so as to predict the underlying mechanisms determining paternity in multiply mated females. The models require: 1) estimations of proportion of offspring sired by the last male to mate (P2), 2) knowledge of the number of sperm transferred by each male, and 3) knowledge of the sperm storage capacity of the female, should this be limited. They will distinguish between “raffles” (sperm mixing without displacement) and sperm displacement mechanisms. The sensitivity of the techniques can be increased by manipulating the number of sperm transferred by each male. Typically, this can be done by manipulating copula duration or number of ejaculations, given a knowledge of the rate of sperm transfer. Data from two contrasting insect species are fitted to the models to demonstrate the techniques. These models are prospective only, and their limitations are discussed. The principal limitation is that we assume that sperm used for fertilization mix randomly in a “fertilization set” immediately prior to fertilization; in reality this may be difficult to identify. When sperm mixing is very rapid, the fertilization set will often be equivalent to the sperm stores, but with slow mixing, the fertilization set may be equivalent to a much more restricted zone and may change with time.

Calling characteristics of parasitized and unparasitized populations of the field cricket Teleogryllus oceanicus

SummaryExamination of three populations of the field cricket Teleogryllus oceanicus revealed the presence of an acoustically-orienting parasitoid fly, Ormia ochracea (Tachinidae), in the population of crickets that has been introduced to the Hawaiian Islands. The cricket is native to Australia and the Pacific, and the fly is native to North America but has also been introduced to Hawaii. Up to 27% of males and 7% of females in Hawaii were infested with fly larvae. Song structure in the parasitized Hawaiian population was distinct from that of the other two groups, with the Hawaiian crickets showing several reduced song parameters. In addition, onset and cessation of calling at dusk and dawn were more abrupt in the Hawaiian population. These results are consistent with selective pressure from the phonotactic flies to decrease risky calling. Silent males were present in all three populations, suggesting that these noncallers may not represent a unique adaptation to the parasitoid.

Nuptial feeding in tettigoniids male costs and the rates of fecundity increase

SummaryA recent debate has centred on the importance of paternal investment for the origin and maintenance of nuptial feeding in insects. Some authors have argued that the rates of nutrient incorporation are likely to be too slow to allow a male to fertilize the eggs that he helps to produce and cannot be considered as paternal investment. Here I report the results of some experiments that show that the positive effects of nutrient donation on female reproduction for one species of spermatophylax producing tettigoniid can be realized within 24 to 48 h. Furthermore, mating was found to induce a refractory period in the female that was longer than the time taken for the incorporation of nutrients and oviposition. Thus, the nurturant male is likely to fertilize the eggs even if last-male sperm precedence is high. The cost of spermatophylax production was manifest as a 5-day recovery period between matings. Thus males and their donations are likely to be limited resources for females resulting in a reversal of the typical sex roles.

Sexual selection and the allometry of earwig forceps

SummaryPositive intraspecific allometry, the tendency for large individuals to have relatively larger morphological traits, is thought to be more likely for secondary sexual traits than naturally selected traits. This is because secondary sexual traits are often used to signal individual quality and positive allometry should arise where the costs and/or benefits of signalling are size dependent. Here we examine the allometric relationships between forceps length, a sexually selected trait and elytra length, a naturally selected trait, in 42 species of earwig. Both forceps and elytra showed positive allometry. However, the degree of allometry was greater for forceps as predicted. If allometry arises due to sexual selection we would predict a greater degree of allometry in species with more exaggerated secondary sexual traits. Across species, the degree of forcep allometry did increase with forcep exaggeration. The relevance of positive allometry to reliable signalling is discussed.

Copula duration and testes size in the yellow dung fly, Scathophaga stercoraria (L.): the effects of diet, body size, and mating history

SummarySeasonal variation in mean hind tibia length and mean testes length is investigated in the yellow dung fly, Scathophaga stercoraria (L.). There is a cycle in mean hind tibia length and mean testes length over a season. The body size curve peaks later than the testes length curve, showing that there is no fixed relationship between the two variables. The causes of variation in testes size and its influence on copula duration are experimentally examined. Increasing the number of Drosophila eaten per day leads to increased mature testes length. Males with larger testes copulate for longer than males with smaller ones, and smaller males copulate for longer than do larger males. While testes shrivel with successive copulas, copula duration remains constant. The more females a male is prepared to copulate with in a day (up to five), the longer he copulates with each. The shrinkage of the testes of males collected throughout a day suggests that males copulate with an average of 4 females per day. The costs of sperm production are thus shown to have a significant influence on the copula duration.

Variability in call structure and pairing success of male field crickets, Gryllus bimaculatus: the effects of age, size and parasite load

The effects of male body size, age and gregarine parasite load on the structure of calls and success at attracting mates were investigated in a natural population of the field cricket, Gryllus bimaculatus. The syllable rate within chirps was negatively related to male size, and the duration of syllables was positively related to size. Older males had more variable syllable rates within their chirps. Size did not influence pairing success. However, paired males were significantly older than solitary calling males. Parasitic infection influenced the pairing success of young males but not old males. These data are discussed in the context of direct and indirect selection for female mating preferences.

Sperm competition or sperm selection: no evidence for female influence over paternity in yellow dung flies Scatophaga stercoraria

Abstract Recent studies of non-random paternity have suggested that sperm selection by females may influence male fertilization success. Here we argue that the problems originally encountered in partitioning variation in non-random mating between male competition and female choice are even more pertinent to interpreting patterns of non-random paternity because of intense sperm competition between males. We describe an experiment with the yellow dung fly, Scatophaga stercoraria, designed to partition variance in the proportion of offspring sired by the second male, P2, between males and females, and to control for sperm competition. Large males were shown to have a higher P2 than small males but P2 was independent of the size of the female’s first mate. This result might suggest an absolute female preference for large males via sperm selection. However, large males have a higher constant rate of sperm transfer and displacement. After controlling for this effect of sperm competition, large males did not achieve higher paternity than small males. We argue that a knowledge of the mechanism of sperm competition is essential so that male effects can be controlled before conclusions are made regarding the influence of sperm selection by females in generating non-random paternity.

Kin recognition and its influence on mating preferences of the field cricket, Gryllus bimaculatus (de Geer)

Abstract Female field crickets, Gryllus bimaculatus , responded differentially towards courting males of different degrees of relatedness. They evaded and even attacked persistently courting full siblings sigificantly more than half-siblings or unrelated males. The latency to mate was therefore greater for full sibling and half-sibling pairs suggesting that females may prefer unrelated males as mates. Female discrimination was also apparent in post-copulatory behaviour. Females were less passive when mated and guarded by full siblings; they attempted to escape more often from guarding full siblings and halfsiblings and spent longer trying to escape than females mated with unrelated males. They were capable of discriminating between males even when they had had no prior experience of conspecifics. Data suggest that the mechanism of recognition in this species involves the learning of self recognition cues as well as those of individuals previously experienced. However, a females own recognition cue may take priority since experience of unrelated males strengthened a females recognition abilities rather than resulting in her treating familiar unrelated males as kin. Cuticular pheromones are suggested as a likely candidate for a recognition cue.

Dimorphisms and fluctuating asymmetry in the forceps of male earwigs

Male dimorphisms are particularly conspicuous examples of the alternative reproductive strategies employed within some species. Such dimorphisms are thought to exist as genetic polymorphisms under ESS conditions, or to be conditional strategies where exogenous conditions determine the adult body plan. Fluctuating asymmetry (FA) is currently considered to be a fitness correlate of significant use in interpreting the functional significance of secondary sexual characteristics. In particular, negative slopes of FA on trait size are thought to arise in traits whose expression is dependent on condition. We measured forceps lengths and asymmetries in 2 island populations of the European earwig Forficula auricularia and Museum specimens of 5 other earwig (Dermaptera, Forficulidae) species from different genera, that appeared to be dimorphic. In a detailed study of Forficula auricularia we found a significant fit to a statistical model for the identification of dimorphisms and, for all species examined, morphs differed in the slope and/or elevation of the allometric relationship between body size and forcep length. Possible determinates of male dimorphisms are suggested from the data. Contrary to expectation, FA was not found to be greater in the minor morphs. Negative relationships between FA and forceps length were absent in both morphs of species examined from museum collections. Of the two island populations of Forficula auricularia, the smaller and more isolated population had higher FA and a negative relationship between FA and forceps length in the major morph. We discuss these patterns in the light of recent theories of FA and honest signalling.

Acoustically‐orienting parasitoids in calling and silent males of the field cricket Teleogryllus oceanicus

Abstract. 1 On three Hawaiian Islands, the introduced Australasian field cricket Teleogryllus oceanicus Le Guillou (Orthoptera: Gryllidae) was found to be attacked by the phonotactic parasitoid tachinid fly, Ormia ochracea Bigot. 2 Noncalling males occurred with callers in all locations, but silent males were more heavily parasitized than callers. 3 Body size was unrelated to both calling status and the likelihood of harbouring parasitoid larvae. 4 An experiment examining the likelihood of calling in the laboratory by males collected as silent or calling individuals showed no difference between the two classes of males, after accounting for parasitoid levels; males harbouring larvae were less likely to call.