Large male advantage for access to females: evidence of male-male competition and female discrimination in a territorial salamander

Summary In the natural habitat visited in this study, adult male red-backed salamanders (Plethodon cinereus) found near females were significantly larger than adult males found alone; there was no evidence for size assortative associations. Previous research has indicated that females associate preferentially with males that occupy high quality territories and that larger individuals are more successful at obtaining higher quality areas. In laboratory experiments, when resource quality was held constant and males were restrained so that male-male interactions were prohibited, females discriminated behaviorally between large and small males by spending more time visually and chemically assessing larger males and more time apparently attempting to leave the chamber when near smaller males. Females were found near the larger males at the end of the trials significantly more often than predicted by chance. In a separate experiment in which only females were restrained, larger males spent significantly more time in aggressive postures when paired with smaller males than when alone with the female. Smaller males spent significantly more time in submissive postures when paired with larger males and more time near the female when alone. Therefore, large body size of males may positively affect both intra- and intersexual interactions and, ultimately, mating success of male P. cinereus.     

Factors leading to the evolution and maintenance of a male ornament in territorial species

Male ornamentation is assumed to have evolved primarily from selection by female mate choice. Yet this is only one possible reason for ornament evolution. Ornaments might also be useful in aggressive competition by improving opponent assessment between males, or they might function to enhance signal detection by making males more conspicuous in the environment. We tested both these ideas in territorial Anolis lizards in which female choice is either absent or secondary to males competing for territories that overlap female home ranges. Male tail crests only evolved in species in which territory neighbors were distant, consistent with the signal detection hypothesis. Once the tail crest had evolved, however, it seems to have become a signal in itself, with variation in the frequency and size of tail crests within species correlating with variables predicted by the aggressive competition hypothesis. Our study presents an apparent example of a male ornament in which the selection pressure leading to variation among species in ornament expression is different from the selection pressure acting on variation within species. The Anolis tail crest is therefore likely to be an exaptation. We caution that conclusions made on the evolution of male ornaments are dependent on the phylogenetic perspective adopted by a study. Studies restricted to single species are useful for identifying selection pressures in contemporary settings (i.e., the current utility of traits), but may lead to erroneous conclusions on the factors that initially lead to the origin of traits.     

Extra-pair paternity,sperm competition and the evolution of testis size in birds

Sperm competition should select for increased sperm production if the probability of fertilization by a specific male is proportional to the relative number of sperm inseminated. A review of the literature generally supports the predicted positive association between sperm production or allocation and various measures of the presumed intensity of sperm competition. However, it is not clear how increased sperm competition is related to extra-pair paternity, and it remains unknown whether certainty of paternity should be associated with relative testis size. Based on a large sample of bird species with information on extra-pair paternity gathered from the literature, we demonstrate that testis mass is related positively to the level of extra-pair paternity, after controlling for body size and phylogeny. Although large testes may be necessary to avoid sperm depletion in species in which males frequently engage in multi-pair copulations, we argue that selection has favoured increased testis mass in situations of more intense sperm competition in order to retaliate against copulations by rival males. The fact the males are not always successful in retaliating against rival ejaculates further suggests that females may largely control the allocation of paternity in birds and that increased sperm production by males may simply be a male strategy to make the best of a bad situation.     

Two roads to two sexes: unifying gamete competition and gamete limitation in a single model of anisogamy evolution

Recent studies have revealed the importance of self-consistency in evolutionary models, particularly in the context of male–female interactions. This has been largely ignored in models of the ancestral divergence of the sexes, i.e., the evolution of anisogamy. Here, we model the evolution of anisogamy in a Fisher-consistent context, explicitly taking into account the number of interacting individuals in a typical reproductive group. We reveal an interaction between the number of adult individuals in the local mating group and the selection pressures responsible for the divergence of the sexes. The same underlying model can produce anisogamy in two different ways. Gamete competition can lead to anisogamy when it is relatively easy for gametes to find each other, but when this is more difficult and gamete competition is absent, gamete limitation can provide another route for anisogamy to evolve. In line with earlier models, organismal complexity favors anisogamy. We argue that the early contributions of Kalmus and Scudo, largely dismissed as group selectionist, are valid under certain conditions. Linking their work with the contributions of Parker helps to explain why precisely males keep producing more sperm than can ever lead to offspring: sperm could evolve to provision zygotes but this brings little profit for the effort required, because sperm would have to be equipped with provisioning ability before it is known which sperm will make it to the fertilization stage. This insight creates a logical link between paternal care under uncertain paternity (where again investment is selected against when some investment never brings about genetic benefits) and gamete size evolution.     

Complex courtship in a bimodal grasshopper hybrid zone

Grasshoppers of the Chorthippus albomarginatus-group, which is outstanding with respect to its complex courtship song, were studied at fifteen localities in the Ukraine and Moldova. The analysis of the courtship songs revealed two species: C. albomarginatus in north-eastern Ukraine and Chorthippus oschei in the south-western Ukraine and Moldova. In a belt about 200 km wide, not only were one or the other pure species found, but also males with intermediate song characters. C. albomarginatus and C. oschei were hybridised in the laboratory, and F1 hybrid males as well as F2 hybrid males produced intermediate song patterns, quite similar to those recorded in the field. We defined a "hybrid song score" for intermediate songs. The score showed a bimodal distribution with most songs resembling one or other parental type, but with only a few intermediates. At several localities, where hybrids with songs similar to one of the parental species dominated, some individual males sang more similarly to the other species. In one locality, two hybrid populations only 3 km apart had different parental types. Hybrid songs can contain novel elements, even more complex than the parental ones, which may offer a new starting point for sexual selection. We suggest that genetic introgression occurs between the two sibling species C. albomarginatus and C. oschei within a wide hybrid zone stretching over a distance of several hundred kilometres, but with a patchy spatial distribution. Electronic Supplementary Material Supplementary material is available for this article if you access the article at . A link in the frame on the left on that page takes you directly to the supplementary material. Communicated by D. Gwynne     

Tardy females,impatient males: protandry and divergent selection on arrival date in the two sexes of the barn swallow

Protandry reflects the earlier arrival of males than females to the site of reproduction. Such protandry is hypothesised to arise from sex differences in costs and benefits of early arrival. I investigated temporal patterns of arrival date of male and female barn swallows Hirundo rustica and temporal patterns of selection to test the hypothesis that sex differences in selection account for sex differences in arrival date. Mean arrival date of male barn swallows but not of females advanced during the last 33 years, giving rise to an increasing sex difference in arrival date. Early arrival was favoured by increasingly better survival in males, while females showed an opposite pattern that did not reach significance, although the effect differed between sexes. Early arrival increased fecundity in both sexes equally.The sex difference in viability selection in relation to arrival date changed from positive to negative as the degree of protandry increased in recent years, although there was no similar significant relationship for fecundity selection. Furthermore, sex differences in viability selection in a given year affected the degree of protandry in the following year through differential survival of certain phenotypes over others. Finally, temporal changes in sex difference in viability selection and protandry were related to an increase in the interval between first and second clutches, as the duration of the breeding season increased because of climatic amelioration. These findings suggest that arrival date is under divergent selection in the two sexes, providing a mechanism for the evolution of protandry.     

Thermoregulatory cost affects territorial behavior in hummingbirds: a model and its application

A common assumption in behavioral ecology is that the valuation of a resource by consumers depends on the energetic value of the resource itself. Nevertheless, the value of a resource may be relative to the condition of the organism, which is in turn related to the abiotic conditions such as ambient temperature. We developed a theoretical model—incorporating these untested assumptions—to predict a functional relationship between territorial aggression and ambient temperature for individuals sensitive to daily variations in energy availability. We evaluated our theoretical predictions against a field experiment carried out with the hummingbird Sephanoides sephaniodes. The model predicted a quadratic relation between aggression intensity and ambient temperature. Field data were better explained by a quadratic equation than a linear function, suggesting the existence of lower and upper thresholds of temperature which determine the intensity of territorial defense. Ambient temperature affects energy expenditure for thermoregulation, and therefore, it fixes the benefit level that must be produced by the territory to pay the costs of its defense. Our findings strongly suggest that abiotic conditions can change an animal evaluation of the yield of a resource and in turn influence the behavioral strategy which it adopts.     

Defensive secretion of a flightless grasshopper: Failure to prevent lizard attack

Summary We tested the role of a grasshopper defensive secretion in deterring lizard predation. Adults, but not young larvae, of the chemically defended lubber grasshopperRomalea guttata (=microptera) froth a volatile secretion when attacked by predators. The lizardAnolis carolinensis failed to strike juvenile lubbers (which lack secretion) in laboratory trials. Survivorship of palatable crickets loaded with secretion offered toA. carolinensis was not significantly different from survivorship of control crickets. In experiments designed to investigate if lizards learn an aversion to the secretion, striking times forSceloporus undulatus fed wax worms coated with secretion were not significantly different over three days of trials. Three primary conclusions are drawn from these data. First, the secretion may not be necessary for lubber protection from lizards. Second, lubber secretion does not appear to deter lizards from attacking or eating prey items. Third, lizards do not appear to develop an aversion to the secretion.     

Patterns of use of space in a guild of territorial reef fishes

There exists on Heron Reef, at the southern end of the Great Barrier Reef, an 8-species guild of ecologically very similar, territorial, herbivorous reef fishes. All individuals of these species maintain territories on rubble substrata throughout juvenile and adult life. Territories are defended from all other guild members. Three rubble patches, each containing residents of 3 guild species: Pomacentrus apicalis, P. wardi, and Abudafduf lachrymatus, have been monitored for 12 to 18 months. This paper examines the patterns of use of space as shown by the antecedent and subsequent histories of sites chosen by 43 new colonists, and sites vacated through the loss of 34 residents on the rubble patches. In addition, variation through time is examined in the total amount of space held in territories on each rubble patch. The 3 species show similar preferences for space as colonists, although adult and juvenile colonists behave differently. Numbers of colonists detected on rubble patches are not proportional to the resident populations of the 3 species. Residents of the 3 species are equal in their abilities to enter spaces vacated through mortality, although they differ slightly in methods used to enter sites. They do not respond preferentially to sites previously occupied by any particular species. P. wardi shows a higher rate of mortality than the other species, and residents of this species are more often dislodged by new colonists. The total amount of space held on any rubble patch did not vary during the year from September, 1972 to October, 1973. The data are interpreted as supporting the hypothesis that the 3 species do not differ in their requirements for space on rubble patches, and that they are in competition for a short supply of such space. Differences exist in their strategies for obtaining and holding such space. These differences are important for explaining the continued presence of P. wardi on rubble patches. P. apicalis and a. lachrymatus are both specialists in holding territories on the upper reef slope. No differences have been detected in their requirements or competitive abilities. P. wardi is a fugitive species on the upper reef slope, coexisting because it maintains a refuge from competition by occupying some un-preferred sites off rubble patches.     

Determinants of ejaculate size in a grasshopper (Chorthippus parallelus)

Models for the evolution of ejaculate expenditure predict that ejaculate size (ES) depends on male body condition, on female fecundity, or on the risk of sperm competition. These ideas were tested in the grasshopper Chorthippus parallelus. ES ranged over four orders of magnitude, around a mean of 442,000 sperm. ES was independent of male body size and female fecundity. None of the male condition parameters - age, degree of parasitism, macropterism score (a measurement of competitive energy allocation to other sources), and the mass-femur length residuals - explained variation in ES. As expected in species where females are cyclically non-receptive after mating, male-male encounters before copulation were a more reliable indicator of the risk of sperm competition than the number of males present during copulation. In particular, the summative number of male-male encounters rather than the male-male encounter frequency was a significant predictor of ES variation (24.9%).     

Local adaptation and sexual selection: a reciprocal transfer experiment with the grasshopper Chorthippus biguttulus

Immigration into locally adapted populations has been suggested, among other potential causes, to maintain genetic variance in fitness necessary for good-genes models. Using a reciprocal transplant experiment we examined whether females prefer native to transferred males in the grasshopper Chorthippus biguttulus. On average, native and transferred males did not differ in their attractiveness, measured as female response rate to playbacks of male acoustic courtship signals. In line with this result, we found no significant effect of transfer on body size, condition, fluctuating asymmetry or song traits. However, the reciprocal transplant experiment showed that environmental conditions did influence body condition and maximum loudness of the calling song, but that the genetic origin of male grasshoppers had no significant effect on any of the analysed traits.Communicated by L. Simmons     

Sexual selection in the northern pintail (Anas acuta): the importance of female choice versus male-male competition in the evolution of sexually-selected traits

We experimentally studied the relative importance of plumage, dominance status, and courtship behavior in determining male pairing success in the northern pintail Anas acuta and assessed whether these traits function in female choice, male-male competition or both. In an experiment (experiment IA) that eliminated the confounding effects of male-male competition and social courtship, females chose males with pure white breasts and colorful scapular feathers. When the same group of birds were free to interact (experiment 1B), male behavior was more important: females chose males that courted them intensely and were attentive to them, although preferred males again had whiter breasts and more colorful scapulars. In a second experiment (experiment 2), testing the effect of age on pairing success, females showed a significant preference for 2-year-old males over yearlings: 2-year-old males courted more and were more attentive to the female than yearlings; they were also more colorful than yearlings in a number of plumage measurements. Although males (in both experiments 1B and 2) were aggressive to one another while courting the female and dominant males were sometimes able to exclude subordinates from social courtship, contrary to expectation, we found no relationship between initial dominance rank and pairing success or dominance rank and age. In addition, dominance was not correlated with any of the morphological traits measured. Once chosen, however, subordinate males typically initiated fights with the higher-ranked male(s) and quickly achieved dominance. These results suggest that (1) females choose males based on a suite of morphological and behavioral characteristics, (2) male dominance relationships do not constrain active female choice, (3) a male's position in a dominance hierarchy is largely a result rather than a cause of female choice, and (4) female choice plays a more significant role than male-male competition in the evolution of several secondary sexual traits in male northern pintails.     

Intrasexual selection in Mirza coquereli : evidence for scramble competition polygyny in a solitary primate

The primates of Madagascar (Lemuriformes) deviate from fundamental predictions of sexual selection theory in that polygynous species lack sexual dimorphism, have even adult sex ratios and often live in female-dominated societies. It has been hypothesized that intrasexual selection in these species is either reduced or primarily focused on traits related to scramble competition. The goal of this study was to examine these hypotheses by studying the mating system of a solitary nocturnal species, Mirzacoquereli. During a 4-year field study in western Madagascar, I captured and followed 88 individually marked animals. I found that adult males were significantly larger than females, providing the first evidence for sexual size dimorphism in lemurs. In addition, the adult sex ratio was biased in favour of females in 3 out of 4 years. There was no significant sex difference in canine size, however. Males showed pronounced seasonal variation in testis size with a 5-fold increase before and during the short annual mating season. During the mating season, males had more injuries than females and more than quadrupled their home ranges, overlapping with those of more than ten females, but also with about the same number of rivals. Only about one social interaction per 10 h of observation was recorded, but none of them were matings. Together, these results indicate that these solitary lemurs are clearly subject to intrasexual selection and that male-male competition is primarily, but not exclusively, of the scramble type. In addition, they suggest that the above-mentioned idiosyncracies may be limited to group-living lemurs, that social systems of solitary primates are more diverse than previously thought, and that the temporal distribution of receptive females is responsible for this particular male mating strategy. Received: 11 January 1997 / Accepted after revision: 18 April 1997     

New birds on the block: new neighbors increase defensive costs for territorial male willow ptarmigan

Previous researchers have hypothesized that site-faithful animals may benefit from the presence of familiar neighbors. This study compares the relative costs of territorial defense against new and former neighbors by male willow ptarmigan (Lagopus lagopus). Territorial defense against new neighbors appeared to require a greater expenditure of both time and effort than did defense against former neighbors. Territorial males that had several new neighbors spent a higher proportion of time fighting than did males with fewer new neighbors, and males with both new and former neighbors spent a greater amount of time fighting with their new neighbors, on average, than with their former neighbors. In addition, fights with new neighbors occurred relatively more frequently and were longer than fights with former neighbors. Finally, fights involving new neighbors tended to escalate to higher levels than fights between former neighbors. Reduced defensive costs for site-faithful, territorial males may provide one explanation for the tendency of males to be more site-faithful than females in many species.     

Rules of engagement for adult salamanders in territorial conflicts with heterospecific juveniles

Coexisting species within a guild have the potential for resource overlap and consequently for interspecific competition (e.g., interspecific territoriality). When the adults are of different sizes, which frequently is the case in terrestrial salamander communities in eastern North America, competition may occur between juveniles of the larger species and adults of the smaller species. Adults of the relatively small redbacked salamander (Plethodon einereus: up to 13 cm total length) defend intra- and intersexual territories on the forest floor, and they are broadly sympatric with the larger P. glutinosus (up to 21 cm total length). Although individuals of the two species share the same forest floor habitat, we found significantly fewer juveniles of P. glutinosus sharing territories with 336 same-size adults of P. cinereus than would be expected by random chance alone. In laboratory experiments, residential adults of P. cinereus were as aggressive toward juvenile intruders of P. glutinosus as they were toward adult conspecific intruders. Therefore, adults of P. cinereus appear to defend territories against juveniles of P. glutinosus, illustrating how interference competition may depend on the symmetry of sizes between the species.     

Symmetric fights as a measure of escalation potential in a symbiotic,territorial snapping shrimp

Summary The shrimp Alpheus armatus territorially defends the sea anemone it occupies, using as a weapon its large, specially modified snapping claw. This behavior was studied in experimental contests which were symmetric (matched individuals) with respect to sex, size, and residency. Characteristics of these contests were compared for two size-classes of male and female shrimp.There were no significant differences between these groups of shrimp in the number of bouts required to establish dominance or in the number of snaps exchanged. Large females had shorter contests than either small or large males, and losers of contests between large females were injured more frequently and more severely.This distinctiveness of large female contests could be interpreted as evidence that (i) controlling anemones is more important for large females, (ii) injuries are less important for large females, or (iii) large females lose the ability to assess one another because their contests are less frequent.If injuries are an accurate measure of the most important costs associated with fighting, then these results indicate that short contests are not necessarily the least costly, and that females can be more aggressive than males, as measured by escalation potential, in sexually selected species.     

Implications of a simple competition model for the stability of an intercropping system

It is a common view that intercropping systems of agricultural crops produce more stable yields than do systems in which the same crops are grown in monoculture. This paper discusses a modelling approach which has been used to support the notion that whether or not intercropping is more stable than monoculture depends on the mode of interaction among crops, i.e. whether two different crops suppress or enhance each other. It is shown here that this notion is not supported by the model used. We conclude that the relative merits of the two cropping systems depend on the proportion of land allocated to each crop rather than on the mode of interaction. The model suggests that if the optimum allocation of land is considered, both systems will be equally stable.     

Resources and offspring provisioning: a test of the Trexler-DeAngelis model for matrotrophy evolution

Theoretical models of the evolution of matrotrophy from a lecithotrophic ancestor suggest that resource availability plays a major role in selective scenarios favoring a change in offspring provisioning. We examined effects of feeding level on embryo provisioning in the livebearing fish Gambusia geiseri, a species with dual provisioning of embryos via both yolk sequestered in large eggs and post-fertilization mother-to-embryo nutrient transfer. Females were fed either once per day or once every three days for three months. Females fed daily had marginally larger brood size, significantly larger embryos, and a higher rate of nutrient transfer (assayed directly by injection of radiolabeled nutrients) than females fed every third day. There was no difference in the size of unfertilized eggs between the feeding treatments. Resource effects on matrotrophic provisioning in G. geiseri suggest that matrotrophy plays an important role in provisioning and allows females to adjust offspring size in response to resource availability.