Food threshold territoriality in a water strider (Gerris remigis)

Summary Summer generation 3rd, 4th and 5th instar nymphs plus adults of Gerris remigis were satiated for 2 days in a laboratory tray then deprived of food. Within 1/2–2 days, 19 of 27 nymphs and 10 of 30 adults began to exhibit territoriality, continued being so for 3–9 days, then ceased shortly before becoming quiescent. In the field, muscid flies fed to different territorial striders at a rate of 0, 1, 2, or 3 flies/day, resulted in 8 of 10 striders (at 2 flies/day) and 10 of 10 striders (at 3 flies/day) ceasing territoriality, whereas 5 of 8 controls (at 0 flies/day) remained territorial. Thus, lower and upper food thresholds were demonstrated, the upper threshold approached both from above (laboratory study) and below (field study).     

Role of female territoriality in social and mating systems of Canthigaster valentini (Pisces: Tetraodontidae): evidence from field experiments

Sharpnose puffers, Canthigaster valentini (Pisces: Tetraodontidae) at Lizard Island, Australia, live in made-dominated haremic social and mating systems. The hypothesis was that mature females are restricted in their movements and can be monopolized by some males. Field experiments at Lizard Island, Great Barrier Reef, between January and March 1983 showed that mature females were still territorial in the absence of males and movements of females were not controlled by males. Males abandoned their territories when their females were removed. The territorial behavior of those males with access to females (territorial males) restricted the access of other males (bachelor males) to them. Bachelor males took over harems and became territorial males when established territorial males were removed. The results of the experiments thus supported the hypothesis.     

Sources of variation in male mating success and female oviposition rate in a nonterritorial dragonfly

Summary Variance in male mating success and female oviposition rate was studied on a lifetime and daily scale in a nonterritorial libellulid dragonfly. While controlling for weather, population density, and sex ratio, we investigated whether phenotypic variation could explain a significant amount of the observed variation. By collecting most of the data in a large outdoor field enclosure, we had access to (1) real lifetime data as emigration was prevented, (2) exact values of density and sex ratio, and (3) phenotypic data that are virtually unavailable under natural conditions. Variance in LMS was higher in males than variance in lifetime number of ovipositions in females. Lifespan was of major importance in explaining the variation. Males readily remated on the same day whenever possible (maximum 6 times), whereas in females each oviposition bout usually was followed by a nonreproductive period of approximately 4 days. An analysis of the sources of variance in daily reproduction rate showed that, besides the expected effects of weather, an excess of one sex resulted in a negative effect on the individual mating probability of members of that sex. Even after correction for weather and population parameters, phenotypic traits explained a significant amount of the remaining variation: males with a high mating rate were large, agonistic, active mate searching, and long-living individuals. Females were more likely to oviposit when they showed higher flight activity and when more days had elapsed since the previous oviposition. The presence of a strong size effect on male mating probability, and its absence in females, was confirmed at a pond (native habitat). We propose a model on the causal relationships between the relevant parameters and the variation in reproductive success. Finally, we briefly discuss why variance in reproductive success is not a very good measure of the opportunity for sexual selection in this species. Offprint requests to: N.K. Michiels at the present address     

A simulation approach to evolutionary game theory: the evolution of time-sharing behaviour in a dragonfly mating system

Summary The time-sharing behaviour of dragonfly males (Aeschna cyanea, Odonata) at the mating place is investigated. By computer simulation of the males' interactions the expected number of copulations for different strategies is quantified. The simulation results show that a male's mating succes depends on his strategy and that a strategy's specific pay-off depends on the frequency distribution of strategies in the population. Based on these results, a method for the simulation of the evolutionary process is presented. This approach to evolutionary game theory allows the prediction of evolutionarily stable strategies (ESS), even when the system studied is far too complex for an analytical solution. For the time-sharing behaviour of Ac. cyanea males at the mating place the ESS predicted by the simulation approach complies with field observations.     

Availability of receptive females at the mating place and mating chances of males in the dragonfly Aeschna cyanea

Summary The females of the dragonfly Aeschna cyanea visiting ponds vary in their receptiveness to males waiting there. Individual males pay several short visits to a mating place in the course of a day, searching for receptive females. Non-receptive females prefer to oviposit at the pond in the evening, whereas receptive females arrive throughout the day. The highest number of receptive females is found on warm days during the afternoon at which time the greatest number of copulations occur. However, as the number of males and, therefore, the competition between males is also highest during these periods, the mating chances of a particular male are constant throughout the day.     

The effects of age and relatedness on mating patterns in thornbug treehoppers: inbreeding avoidance or inbreeding tolerance?

The social environment of many species includes synchronous maturation of siblings in family groups, followed by limited dispersal of adults from their natal site. Under these conditions, females may experience high encounter rates with same-age siblings during mate searching, increasing their risk of inbreeding. If inbreeding depression occurs, mating with a sibling is often considered maladaptive; however, in some contexts, the inclusive fitness benefits of inbreeding may outweigh the costs, favoring females that tolerate some level of inbreeding depression. We evaluated mating patterns in the treehopper Umbonia crassicornis, a semelparous species in which females encounter same-age siblings during mate searching. A female U. crassicornis that mates with a brother suffers from inbreeding depression. We used a free-choice mating design that offered females simultaneous mating opportunities with three groups of males: siblings, same-age nonsiblings, and older nonsiblings. These groups represent the types of males typically encountered by females during mate searching. Our goal was to assess whether mating patterns were influenced by inbreeding avoidance by evaluating two hypotheses: kin discrimination and age-based mating (older males cannot be siblings in this species). There was no difference in the proportions of females mating with siblings vs nonsiblings, suggesting an absence of kin discrimination. However, females mated with a greater proportion of older vs younger males. Given that females do not avoid siblings as mates despite a cost to inbreeding, our results provide a possible example of inbreeding tolerance. We also discuss some factors that may have contributed to the mating advantage of older males.     

The mating system of Hanuman langurs: a problem in optimal foraging

Using data from 23 populations of Indian langurs (Presbytis entellus), we show that the proportion of one-male groups is a function of the males’ ability to defend groups of females, as predicted by the monopolisation model of Emlen and Oring (1977). However, we show that in addition to the number of females in the group and their reproductive synchrony, home range size is also a factor, probably because it affects a male’s willingness to search for other groups. These variables in turn are shown to be functions of environmental parameters (rainfall volume and seasonality, and mean ambient temperature) which probably act via the density of resources and refuges in habitats. Received: 3 March 1994 / Accepted after revision: 24 June 1996     

The genetic mating system of a sea spider with male-biased sexual size dimorphism: evidence for paternity skew despite random mating success

Male-biased size dimorphism is usually expected to evolve in taxa with intense male–male competition for mates, and it is hence associated with high variances in male mating success. Most species of pycnogonid sea spiders exhibit female-biased size dimorphism, and are notable among arthropods for having exclusive male parental care of embryos. Relatively little, however, is known about their natural history, breeding ecology, and mating systems. Here we first show that Ammothella biunguiculata, a small intertidal sea spider, exhibits male-biased size dimorphism. Moreover, we combine genetic parentage analysis with quantitative measures of sexual selection to show that male body size does not appear to be under directional selection. Simulations of random mating revealed that mate acquisition in this species is largely driven by chance factors, although actual paternity success is likely non-randomly distributed. Finally, the opportunity for sexual selection (I _s), an indirect metric for the potential strength of sexual selection, in A. biunguiculata males was less than half of that estimated in a sea spider with female-biased size dimorphism, suggesting the direction of size dimorphism may not be a reliable predictor of the intensity of sexual selection in this group. We highlight the suitability of pycnogonids as model systems for addressing questions relating parental investment and sexual selection, as well as the current lack of basic information on their natural history and breeding ecology.     

Evolution of self-organized division of labor in a response threshold model

Division of labor in social insects is determinant to their ecological success. Recent models emphasize that division of labor is an emergent property of the interactions among nestmates obeying to simple behavioral rules. However, the role of evolution in shaping these rules has been largely neglected. Here, we investigate a model that integrates the perspectives of self-organization and evolution. Our point of departure is the response threshold model, where we allow thresholds to evolve. We ask whether the thresholds will evolve to a state where division of labor emerges in a form that fits the needs of the colony. We find that division of labor can indeed evolve through the evolutionary branching of thresholds, leading to workers that differ in their tendency to take on a given task. However, the conditions under which division of labor evolves depend on the strength of selection on the two fitness components considered: amount of work performed and on worker distribution over tasks. When selection is strongest on the amount of work performed, division of labor evolves if switching tasks is costly. When selection is strongest on worker distribution, division of labor is less likely to evolve. Furthermore, we show that a biased distribution (like 3:1) of workers over tasks is not easily achievable by a threshold mechanism, even under strong selection. Contrary to expectation, multiple matings of colony foundresses impede the evolution of specialization. Overall, our model sheds light on the importance of considering the interaction between specific mechanisms and ecological requirements to better understand the evolutionary scenarios that lead to division of labor in complex systems. ELECTRONIC SUPPLEMENTARY MATERIAL: The online version of this article (doi:10.1007/s00265-012-1343-2) contains supplementary material, which is available to authorized users.     

Costs of courtship and mating in a sexually cannibalistic orb-web spider: female mating strategies and their consequences for males

The costs of courtship and mating may include increased risks of predation, the transmission of pathogens, and a loss of foraging opportunities. Thus, a female's decision to tolerate a courting male will depend upon how these costs offset the benefits of mating, which will depend on her reproductive and nutritional status. While these costs may be similar for mated and unmated females, the benefits of mating will be less for mated than virgin females. However, the cost of lost foraging opportunities may be higher for females with fewer nutritional reserves necessary for forming eggs. We examined how these costs and benefits influence the courtship and mating behaviour of male and female orb-web spiders, Argiope keyserlingi. In the field, females on webs that also contained a courting male intercepted fewer prey items per hour than females on webs without a male. In the laboratory, the presence of a courting male at the hub also attracted mantid predators to the web, increasing the risk of predation for both male and female. Staged mating experiments in the laboratory revealed that the frequency of female attacks and pre-copulatory cannibalism was greater among mated than virgin females. Feeding history did not affect aggression in virgin females but, among mated females, food-deprived spiders attacked and cannibalized males more frequently than sated females and only the latter ever remated. These differences in female behaviour influenced male mating strategies. Choice experiments demonstrated that males preferred to venture onto the silk threads of virgin rather than those of mated females. Similar patterns of mate selectivity were observed in the field; females with narrow abdomens attracted more males to the webs than females with broad abdomens, and copulations were observed more frequently among females with narrow abdomens. These smaller females are likely to be virgins that have recently molted. Males that preferentially mate with virgin females will not only avoid potentially fatal attacks but also obtain, on average, a higher fertilization success.     

Why insects swarm: testing the models for lek mating systems on swarming Empis borealis females

Summary Female dance-flies, Empis borealis L., gather to swarm, and males carrying nuptial gifts visit swarms for mating. Field observations and experiments were performed on this behaviorally sex-role reversed species to test models of lekking behavior. The key predictions were: (1) female preference model: male visiting rate and mating rate should increase with the number of females in swarm (swarm size), (2) hotspot model: male visiting rate should be independent of swarm size, and (3) hotshot model: swarm size should be positively correlated with the body size of the largest female in swarm. We found that male visiting rate and mating rate increased with swarm size, and that mating rate per female increased with swarm size. Males also mated more often in larger swarms than in smaller ones. Both males and females visited swarm sites even in the absence of other individuals. When females were successively removed from swarm sites more males than females on average arrived at these sites: 2.25 males per female. When no individuals were present at the swarm site, arriving males moved on to another site, whereas arriving females generally stayed. Larger experimental swarm-markers attracted both more males and more females and even more males when swarming females were present. There was no correlation between mean or median female size in swarms and the number of females in swarms. Thus, the female preference model and the hotspot model were corroborated, while other models were judged unlikely to explain swarming behavior in E. borealis. Correspondence to: B.G. Svensson     

Incorporating Lindenmayer systems for architectural development in a functional-structural tree model

LIGNUM is a functional-structural tree model that represents coniferous and broad-leaved trees with modelling units corresponding to the real structure of trees. The units are tree segments, axes, branching points and buds. Metabolic processes are explicitly related to the structural units in which they take place.This paper enhances the modelling capabilities of LIGNUM with the possibility to formally describe the architectural development of trees with Lindenmayer systems. This is achieved by presenting an algorithm to convert tree structures generated by Lindenmayer systems to the LIGNUM representation of trees with feedback of results of events or processes from LIGNUM to Lindenmayer system. We then give two example applications that model the development of Scots pine (Pinus sylvestris L.) and the dwarf shrub bearberry (Arctostaphylos uva-ursi L.). Finally we discuss our approach and its consequences for the future development of LIGNUM.     

Sex-specific conditional mating preferences in a cichlid fish: implications for sexual conflict

Conditional mating strategies enable individuals to modulate their mating behaviour depending on ‘individual status’ to maximise fitness. Theory predicts that variation in individual quality can lead to differences in mating preferences. However, empirical evidence is scarce particular in terms of variation in male and female strategies. Here, we experimentally investigated quality-dependent variation in mating preferences concerning reliable quality indicators in Pelvicachromis taeniatus, a colourful cichlid fish with mutual mate choice and ornamentation. Males as well as females were artificially manipulated in phenotypic quality by different feeding regimes. Ornamentation was connected to individual quality in both sexes. Males and females showed conditional mating strategies in different directions. Males showed prudent choice by preferring females of similar quality. In contrast to males, low-quality females preferred highly ornamented males, whereas high-quality females showed neither preferences for high- nor for low-quality males. The results suggest that individuals aim for specific benefits depending on individual quality. Furthermore, the conflicting conditional mating preferences of males and females might lead to sexual conflict, implicating a highly dynamical mating system that evolves even in absence of environmental changes.     

Intraspecific brood adoption in convict cichlids : a mutual benefit

Summary Convict cichlids (Cichlasoma nigrofasciatum) are biparental, substrate brooding cichlid fish which have extended care of their eggs, larvae (wrigglers) and free-swimming young (fry). Field observations indicate that intraspecific brood adoption of fry occurs frequently under natural conditions. Of 232 broods 29% contained foreign fry and of 232 broods 15% were identified as fry donors. Foreign fry were usually of similar size to or smaller than the host brood fry. Experimental fry transfers showed that parents accept foreign fry smaller than their own but immediately reject or eat foreign fry larger than their own. A predation experiment showed that within a mixed brood, smaller fry are eaten more often than larger fry. Thus, accepting foreign fry may benefit parents through the dilution effect and through a differential predation effect. In comparison with intact families, broods from which the male parent was removed were less likely to reach independence, had fewer fry surviving to independence, and had a greater probability of having some fry transferred to neighbouring broods. We propose that intraspecific brood adoption in convict cichlids is mutually beneficial to the adult donors and recipients. Correspondence to: B.D. Wisenden     

Evolution of mating systems and sexual size dimorphism in North American cyprinids

Mating systems evolve with sexual size dimorphism (SSD) in many animals. Mating systems with males larger than females occur when males compete for female access or guard territories, while mating systems with group mating tend to occur in species where females are the same size or larger than males. In addition to variation in SSD with mating system, sperm competition varies among mating systems in predictable patterns. We examined the evolution of mating systems with SSD and testes mass in 111 North American Cyprinidae fishes using phylogenetic comparative methods. Our results demonstrate that the evolution of mating systems in Cyprinidae fishes is from ancestral taxa that are group spawners with females the same size or larger than males to pair spawning systems where males tend to be larger than females. We used an additive model to predict male and female body size from testes mass and mating system. Only mating system varied predictably with SSD. Our results for analyses of hyperallometry (Rensch’s rule) were that individual species of Cyprinidae can have hyperallometry for SSD, but the pattern is not present across all taxa.     

The development of a multi-species algal ecodynamic model for urban surface water systems and its application

An ecodynamic model that can simulate four phytoplankton species has been developed to deal with the unique characteristics of urban river systems which has manmade river profile, flow controlled by gates, severe eutrophication status, and fragile aquatic ecosystem. The ecodynamic model was developed referencing two typical models: the water quality simulation model WASP and ecological model CAEDYM. The model can simulate 11 state variables: dissolved oxygen, carbonaceous biochemical oxygen demand, ammonia nitrogen, nitrate nitrogen, organic nitrogen, inorganic phosphorus, organic phosphorus and four phytoplankton species with zooplankton as a boundary condition. The ecodynamic model was applied to Sihai section of the Beijing urban river system, where serious algal blooms broke out in recent years. The dominant phytoplankton species are Cyanophyta, Chlorophyta, Bacillariophyta, and Cryptophyta. Site-specific data on geometry, meteorology, pollution sources, and existing ecosystem parameters were collected and used for model calibration and verification The model results mimic observed trends of water quality and phytoplankton species succession and can be used for forecasting algal blooms as well as assessment of river management measures.     

The genetic mating system of a sex-role-reversed pipefish (Syngnathus typhle): a molecular inquiry

In the pipefish Syngnathus typhle as in other species of Syngnathidae, developing embryos are reared on the male's ventral surface. Although much laboratory research has been directed toward understanding sexual selection in this sex-role-reversed species, few studies have addressed the mating behavior of S. typhle in the wild, and none has capitalized upon the power of molecular genetic assays. Here we present the first direct assessment of the genetic mating system of S. typhle in nature. Novel microsatellite loci were cloned and characterized from this species, and employed to assay entire broods from 30 pregnant, field-captured males. Genetic analysis of 1340 embryos revealed that 1–6 females (mean = 3.1) contributed to each brooded clutch, the highest rate of multiple maternity yet documented in any pipefish. Evidence of multiple mating by females was also detected. Thus, this population of S. typhle displays a polygynandrous mating system, a finding consistent with previous field and laboratory observations. Our results, considered together with similar studies of other syngnathid species, provide preliminary support for the hypothesis that the genetic mating system is related to the evolution of sexual dimorphism in the fish family Syngnathidae. Received: 19 January 1999 / Received in revised form: 15 April 1999 / Accepted: 9 May 1999