Endocrinology of sociality: Comparisons between sociable and solitary individuals within the same population of African striped mice

The social organization of species ranges from solitary-living to complex social groups. While the evolutionary reasons of group-living are well studied, the physiological mechanisms underlying alternative social systems are poorly understood. By studying group-living and solitary individuals of the same species, we can determine hormonal correlates of sociality without the problem of confounding phylogenetic factors. The African striped mouse (Rhabdomys pumilio) is a socially flexible species, which can be solitary or alternatively form complex family groups, depending on population density and the extent of reproductive competition. We predicted group-living striped mice to show signs of reproductive suppression and social stress, resulting in higher corticosterone but lower testosterone levels when compared to solitary-living individuals. To determine whether differences in social organization correlated with hormonal differences, we collected blood samples from free-living striped mice during four breeding seasons when we experimentally induced solitary-living in philopatric individuals by locally reducing population density. Striped mice that were group-living did not change their corticosterone or estosterone levels during the study, indicating that there was no temporal effect during the breeding season. Striped mice of both sexes had significantly lower corticosterone levels after switching from group- to solitary-living. Solitary males – but not solitary females – had higher testosterone levels than group-living conspecifics. Our results suggest that group-living results in physiological stress and can induce reproductive suppression, at least in philopatric males. The switch to solitary-living may thus be a tactic to avoid reproductive competition within groups, and is associated with decreased stress hormone levels and onset of independent reproduction.     

Temporal and spatial variation in dispersal in the greater flamingo (Phoenicopterus ruber roseus)

We studied movements of individually marked greater flamingos (Phoenicopterus ruber roseus) born in the Camargue, southern France, between their two most important breeding colonies in the western Mediterranean (Camargue and Fuente de Piedra, Spain) from 1986 to 1992. The two sites differ in the frequency with which they offer suitable conditions for breeding. Flamingos have bred each year in the Camargue since 1974, but in only 12 of the past 22 years at Fuente de Piedra. Higher colony fidelity is thus expected in the less variable environment (Camargue), but if dispersal occurs competition might be an important factor causing this dispersal. Following years during which breeding birds in the Camrgue were disturbed (1988 and 1990) a higher proportion of adults changed colonies between breeding attempts (= breeding dispersal, 12.4%), while only 0.4% of flamingos breeding in the Camargue dispersed in the other years. As expected, flamingos breeding at Fuente de Piedra showed a higher rate of breeding dispersal (8.14%). No differences were observed between males and females. The importance of breeding failure as a factor causing breeding dispersal in flamingos was also confirmed by the movements of individual birds. The proportion of young flamingos that moved from their natal colony to start breeding at Fuente de Piedra (= natal dispersal) was independent of sex and age, but increased when breeding access to the Camargue colony was more difficult. However, natal dispersal was also higher in 1988 and 1990 (40.5%) than in the remaining years (1.2%), as was breeding dispersal. We discuss possible ways in which the increased natal dispersal among inexperienced birds could be linked with the increased breeding dispersal of adults in the same year.     

Ectoparasitism as a possible cost of social life: a comparative analysis using Australian passerines (Passeriformes)

The hypothesis that cooperative breeding entails a cost in terms of transmission of ectoparasites was tested by a comparative analysis among sympatric Australian passerines. The general trend found using the allometry method and outgroup analysis indicates that contagious ectoparasites are not more common on cooperatively breeding than on non-cooperatively breeding hosts. Body weight, migratory patterns and relative abundance of hosts are factors far more important than cooperative breeding that affect the levels of ectoparasitism in the host genera studies. Ectoparasitism increases with host body weight and relative adundance, while sedentary host genera tended to show less hippoboscid fly diversity than migratory host genera. There is an interaction between breeding system and migratory pattern when relative density of contagious ectoparasites (i.e. mites, ticks and bird lice) is considered: the number of contagious ectoparasites per host is larger on cooperatively breeding host genera than on non-cooperatively breeding host genera among sedentary passerines, but the trend is reversed for migrant passerines.     

Using recruitment age, territorial fidelity and dispersal as decisive tools in the conservation and management of peregrine falcon ( Falco peregrinus ) populations: the case of a healthy population in Northern Spain

Methods of evaluating population trends have recently received particular attention because of perceived declines in several species during the twentieth century. We have studied demographic traits of the peregrine falcon (Falco peregrinus) population in Bizkaia (Northern Spain) for 11 years. This species suffered a severe decline in the 1950s and 1960s and started to recover in the 1980s and 1990s, although the recovery trends differed between areas. In our study area the peregrine falcon density is one of the highest found in Spain. The frequency of juvenile breeders was 2.0%, the mean age at first breeding being 3.7 calendar years for males and 4.0 cy for females. The territorial fidelity was at least 3.4 years for males and 3.7 years for females. Females dispersed on average 80.5 km and males 51.8 km during their pre-breeding movements, whilst distance between birthplace and breeding territories was on average 108.5 km for females and 64.5 km for males. We studied the relationships between adult turnover, recruitment age, territorial fidelity and dispersal in a healthy population in order to establish population dynamics. A combination of these parameters, not just age at first breeding, could be used as a potential early warning signal indicating future changes, prompting their consequences to be evaluated. This approach could lead to the reclassification of the large-scale health of a population. Its utilization would allow resources to be directed into helping “ailing” subpopulations, detecting causes of decline, and developing adequate recovery strategies.     

Resources and dispersal as factors limiting a population of the tussock moth (Orgyia vetusta), a flightless defoliator

The western tussock moth (Orgyia vetusta) is very abundant on one stand of bush lupine (Lupinus arboreus) at the Bodega Marine Reserve (Sonoma Co, Calif., USA), but is sparse or absent on nearby stands. To determine what controls its abundance, both within the outbreak area and more globally, I performed experimental manipulations of resource availability and dispersal. To measure resource limitation, I inoculated 30 caged and 30 uncaged bushes with a realistic range of numbers of tussock moth eggs. On caged bushes, starvation led to density-dependent reductions in survival, pupal weight and fecundity. Larvae on uncaged bushes experienced density-independent ant predation on early instars and density-dependent emigration by late instars. From the results of this experiment, I predicted the density of a resource-limited tussock moth population. The predictions agreed fairly well with data from the outbreak area in 1992. To measure dispersal by the moth, which has flightless adult females, I released 30,000 tussock moth eggs at a central point in each of two uninfested lupine stands, and censused larvae weekly in a circle of radius 16 m until pupation. Median displacement over one entire generation was only 2 m, or about 2 bush radii. Rearing experiments indicated that bushes outside the out-break area are fully nutritionally suitable for the moth. I conclude that two major factors limiting the population are resources (within the outbreak area) and inefficient dispersal (more globally).     

Age specificity of inbreeding depression during a life cycle ofCallosobruchus chinensis (Coleoptera: Bruchidae)

Age-specific effects of inbreeding on fecundity were assayed for adzuki bean weevil Callosobruchus chinensis by comparing inbred lines and their cross. Four consecutive full-sib matings reduced only 10.3 percent in total fecundity, and did not decrease early fecundity at all until third day from the onset of reproduction. It is suggested that recessive detrimental genes have been eliminated from the early period of adult life span when reproductive value is high. There was a slight tendency that inbreeding depression increased as age proceeded though not statistically significant.     

Emergence and dispersal relative to natal nest in the digger wasp Stizus continuus (Hymenoptera: Crabronidae)

The position of the emerging point has rarely been investigated as a factor possibly affecting the future nest settlement behaviour in Hymenoptera, in particular within nest aggregations. We studied the emergence and dispersion patterns of the digger wasp Stizus continuus. Individuals emerged daily in clumped patterns, possibly revealing a certain synchrony of emergence from the same nests, and protandry appeared both at seasonal and daily level. Differences between the number of females that nested relatively close or far from their emergence holes (EH) were either significant or not, depending on the year, and observed dispersal distances from the natal nests did not differ from those obtained by random simulations. By contrast, females nested close to the nearest conspecific nest. Size did not affect the dispersion patterns. EH are thus not important cues for nest establishment, and conspecific nests are probably the key cue for nest-founding females. In addition, males did not prefer to establish territories close to their natal nest.     

Social and ecological influences on dispersal and philopatry in the plateau pika (Ochotona curzoniae)

Benefits and costs of dispersal and philopatry of the socialplateau pika (Ochotona curzoniae) were studied on the Tibetanplateau for 3 years. Although short-lived, plateau pikas livein cohesive family groups that occupy burrow systems in sedgemeadow habitat Most (57.8%) plateau pikas were philopatric,and dispersal movements were extremely restricted. No juvenilefemales or adult pikas moved more than two family ranges betweenyears; the greatest observed dispersal distances were by twojuvenile males that moved five family ranges from the familyof their birth. Traversing unfamiliar habitat was not a costof pika dispersal because most dispersers settled in familiesthat they could easily visit before dispersal. Dispersal movementsappeared to result in equalization of density among pika families,an expected result if competition for environmental resourcesinfluenced dispersal. Males did not disperse to gain advantagesin competition for mates, as evidenced by their moving to familieswith significantly fewer females. Females, however, moved tofamilies with significantly more males. Males provide abundantpaternal care, and significantly more offspring per female survivedto become adults from families with more adult males per adultfemale. Evidence concerning the influence of inbreeding avoidanceon natal dispersal was indirect. Some males exhibited natalphilopatry; thus some families had opportunity for dose inbreeding.Males and females that dispersed had no opposite-sex relativesin their new families. Philopatric pikas may have benefitedby remaining in families that exhibited low local densities,and philopatric females might have benefited from social cooperationwith relatives.     

The consequences of flooding for the dispersal and fate of poeciliid fish in an intermittent tropical stream

Summary We studied the rainy season dispersal of the fish Poecilia gillii (Poeciliidae) from pools in a steepgradient, intermittent stream in Santa Rosa National Park, Costa Rica. The stream consisted of about 20 pools separated by dry streambed except during two floods and subsequent brief periods of flow. Individually recognizable tags permitted mark-recapture estimates of population size and information on individual movements. The first flood was very severe, with pools losing an average of 75% of their populations (range 12–99%). Most of the lost fish died by becoming trapped in desiccating pools. Males and juveniles were more likely to be lost than were females. Population loss was negatively related to pool volume and positively related to streambed slope. In addition, population loss was positively related to preflood population size when the effects of pool volume and slope were removed indicating that pools with higher densities lost more fish. Of the fish recaptured after the flood, the average proportion found in the pool in which they had been tagged varied from 16%–96%, depending on the area of the stream. Of fish that moved, 92% went downstream. The second flood was less severe though stream flow lasted as long, and there was little effect on the pool populations. Involuntary flushing during the flood and voluntary departure apparently interacted to produce the observed patterns.     

Effects of biparental inbreeding on the evolution of gynodioecy: A model and a case study inChionographis japonica var.kurohimensis

To examine the evolution of gynodioecy under biparental inbreeding, we developed a new theoretical model and tested it by estimating paramaeters incorporated in the model in a gynodioecious population ofChionographis japonica var.kurohimensis. The model and the results showed biparental inbreeding affects the evolution of gynodioecy, but the effect is very small.     

Global phylogeography of the dolphinfish (Coryphaena hippurus): the influence of large effective population size and recent dispersal on the divergence of a marine pelagic cosmopolitan species

Pelagic fish that are distributed circumtropically are characterised by a low population structure level as a result of a high capacity for dispersion and large population sizes. Nevertheless, historical and contemporary processes, including past demographic and/or range expansions, secondary contact, dispersal, gene flow, and the achievement of large effective population sizes, may play a part in the detection of divergence signals, especially in the case of tropical pelagic species, whose distribution range depends strongly on the sea surface temperature. The connectivity and historical demography of Atlantic, Indian, Pacific and Mediterranean populations of dolphinfish (Coryphaena hippurus) was studied using partial sequences of the mitochondrial DNA NADH dehydrogenase subunit 1 (ND1). AMOVA analyses revealed significant inter-oceanic divergence with three phylogroups located in the Indo-Pacific, Eastern Atlantic, and Mediterranean Sea, the last one being the most divergent. However, it was not possible to clearly observe any genetic differentiation between the Indo-Pacific and Atlantic populations, as has been reported for most tropical pelagic species of tuna and billfishes. This supports the assumption of recent dispersal among basins facilitated by the actual continuous distribution of dolphinfish populations. Moreover, the lack of a divergence signal for populations separated by the Panamanian Isthmus reveals that genetic drift does not exert a strong influence on tropical pelagic species with large effective population sizes.     

Potential vs. realized larval dispersal: fish predation on larvae of the ascidian Lissoclinum patella (Gottschaldt)

The total swimming time of an invertebrate larva in a closed container is seldom a reliable indication of its swimming time in the field. In this paper we compare in situ observations on larval dispersal of the colonial ascidian Lissoclinum patella (Gottschaldt) to the total swimming time of the larvae in closed containers. A total of 123 out of 133 larvae were followed from release until they were consumed by predators (87%) or settled (13 %). More than three-quarters of the predation was from pomacentrid fishes [primarily Pomacentrus popei Jordan & Scale and Glyphidodontops rollandi (Whitley)]; the rest was from corals and zoanthids. In closed containers the larvae swam for > 2 h, but larvae followed in situ persisted for < 10 min. The behavior of the larva and the effect of predators greatly reduce the larva's realized dispersal, and may even limit the vertical distribution of adult L. patella. Although L. patella larvae have a potential dispersal distance of several hundred meters (depending upon local current speed), their realized dispersal within a coral reef appears to be < 10 m, and the majority of the larvae end up in fish guts.     

Sorting out the effects of Wolbachia, genotype and inbreeding on life-history traits of a spider mite

Wolbachia bacteria manipulate host reproduction by inducing cytoplasmic incompatibility (CI) and sex ratio distortion. Wolbachia are transmitted from mother to offspring through the cytoplasm of the egg. Therefore, reproduction of Wolbachia is tightly coupled to reproduction of its host. Mathematical analysis predicts that in the course of evolution, traits that reduce the physiological costs of the infection will be selectively favored. For a Wolbachia-host system to evolve, traits under selection must have some genetic component and variation must be present in the population. We have previously established that highly inbred isofemale lines of the two-spotted spider mite Tetranychus urticae may differ regarding the effects of infection by Wolbachia, and that at least some of the traits affected had a genetic component. However, the effects measured could have been affected by the fact that the lines were severely inbred prior to the experiments. In this paper we attempt to distinguish between the effects of Wolbachia, isofemale line, and inbreeding. We show that Wolbachia did not affect longevity but infected females produced smaller clutch sizes, more daughter-biased sex ratios and had decreased F1 mortality; between-line variation was found for clutch size, F1 mortality and sex ratio; finally, inbreeding resulted in an overall reduction of clutch sizes, and a change in survival curves and mean longevity. This revised version was published online in July 2006 with corrections to the Cover Date.     

Consequences of inbreeding for the cowpea seed beetle, Callosobruchus maculatus (F.)(Coleoptera: Bruchidae)

Inbreeding is said to reduce vigour and fitness. It may also determine how a population responds to selection. Local populations of Callosobruchus maculatus , the cowpea seed beetle, are established annually from small numbers of founders and the species has been distributed to many parts of the world where isolated populations may have been founded by very small numbers of individuals. After more than 20 generations of inbreeding, inbred lines have been shown to diverge from a common ancestral stock in similar directions with respect of some variables such as developmental speed, but haphazardly in respect of other parameters such as male weight. The respective roles of drift and of selection as effective evolutionary forces in inbred lines are discussed in the light of these results. It is argued that some intraspecific differences in C. maculatus may be explained as a product of periodic inbreeding, but that the process does not impair the ability to adapt to local conditions so contributing to the status of the species as a pest of international importance.     

Vertical transmission of Nosema fumiferanae (Microsporidia: Nosematidae) and consequences for distribution, post-diapause emergence and dispersal of second-instar larvae of the spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae)

We examined vertical transmission of Nosema fumiferanae in the eastern spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae), and how it affects overwintering distribution and survival and spring emergence and dispersal of second-instar larvae in outbreak populations. Females containing 5.0 x 10(5) spores or more consistently produced 100% infected progeny. Transmission efficiency was still 50% at burdens as low as 0.2 x 10(5) spores per moth. Infection intensity in offspring increased with maternal spore load but became highly variable above 25 x 10(5) spores per female. Nosema multiplied in second instars for at least 1 month after they entered dormancy, regardless of temperature (2 degrees C versus 21 degrees C). Infection did not affect the distribution of overwintering larvae in a white spruce canopy. Dormancy survival between late-summer and the following spring was lower in families from infected females and was negatively correlated with larval infection intensity. Infection delayed larval emergence from hibernacula in the spring and resulted in delayed dispersal of emerged larvae, at least when parasite prevalence and infection intensities were high. Infected larvae were less successful in establishing feeding sites after dispersal. Our results underscore the potential of Nosema infection to negatively affect processes early in the budworm life cycle.     

Short-distance dispersal behavior and establishment of the parasitoid Gonatocerus ashmeadi (Hymenoptera: Mymaridae) in Tahiti: Implications for its use as a biological control agent against Homalodisca vitripennis (Hemiptera: Cicadellidae)

The egg parasitoid Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae), was introduced into French Polynesia as a biological control agent to control the invasive plant feeding pest Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae). The short-distance dispersal of G. ashmeadi was monitored as part of the biological control program. G. ashmeadi showed exponential dispersal capacity with 47 m/day being a minimum estimate of its natural rate of spread at high host densities (>150 nymphs per minute of sweep net sampling) in urbanized areas at sea level, which were characterized by a high diversity of exotic ornamental plants. This rate of spread contrasted starkly with almost nonexistent establishment and dispersal where host densities were very low (<2 nymphs per minute of sweep net sampling) at high elevation (800 m) with relatively undisturbed native vegetation. Survey results across different altitudes revealed an effect of vegetative diversity and host density on the measurable mobility and establishment of G. ashmeadi. In contrast, no significant influence of wind direction was found on G. ashmeadi dispersal rate or direction. Survey results for G. ashmeadi from French Polynesia suggest that the best release establishment strategies for classical biological control of H. vitripennis are: (1) many small releases where host density is high, or (2) larger and fewer releases where host densities are low.     

Morph-dependent fatty acid oxidation in a wing-polymorphic cricket: implications for the trade-off between dispersal and reproduction

Although a considerable amount of information is available on the ecology and physiology of wing polymorphism, much less is known about the biochemical-genetic basis of morph specialization for dispersal versus reproduction. Previous studies have shown that the dispersing morph of the wing-polymorphic cricket, Gryllus firmus, prioritizes the accumulation of triglyceride flight fuel over ovarian growth, while the opposite occurs in the flightless morph during the first week of adulthood. In this study, we compared the in vivo rate of lipid oxidation between genetic stocks of flight-capable versus flightless morphs to determine the role of lipid catabolism in morph specialization for flight versus reproduction. During the first five days of adulthood, in the absence of flight, fatty acid oxidation was substantially lower in the dispersing morph relative to the flightless morph, when either radiolabeled acetate or palmitate was used as a substrate. Differences between the morphs in fatty acid oxidation were genetically based, occurred co-incident with morph-specific differences in triglyceride accumulation and ovarian growth, and were observed on a variety of diets. A genetically based trade-off in the relative conversion of palmitate into CO(2) versus triglyceride was observed in morphs of G. firmus. Decreased oxidation of fatty acid and increased biosynthesis of triglyceride, both appear to play an important role in flight fuel accumulation, and hence morph specialization for flight. Conversely, increased oxidation of fatty acid likely fuels the enhanced ovarian growth in the flightless morph. The results of the present study on fatty acid catabolism, and previous studies on triglyceride and phospholipid biosynthesis, provide the first direct evidence that genetically based differences in in vivo flux through pathways of intermediary metabolism underlie a trade-off between flight capability and reproduction--a trade-off of central importance in insects.     

Mother's timing and duration of corticosterone exposure modulate offspring size and natal dispersal in the common lizard (Lacerta vivipara)

Although multiple condition dependence in dispersal is common, the proximate mechanisms that integrate information from multiple sources remain largely unknown. In the common lizard (Lacerta vivipara), earlier studies have shown that maternal plasma corticosterone level interacts with maternal phenotype to affect offspring phenotype and dispersal strategy, and may reflect the mother's external and/or internal environment. However, the mechanism by which these two types of environmental information are integrated has not been identified. We explored the possibility that the timing and duration of the corticosterone signal are the key factors by which the message is modulated. We found that the timing of corticosterone exposure affects the juvenile phenotype: an exposure to corticosterone early in development has negative effects on juvenile size, weight, and body condition, that can nevertheless be restored in the case of a prolonged exposure. The duration of corticosterone exposure affects the dispersal strategy, although the precise effect depends on the sex ratio of the clutch. That is, in female-biased clutches, a prolonged exposure during gestation promotes philopatry of juveniles, while a short exposure enhances their dispersal, a result that is consistent with kin competition theory. Therefore, our results demonstrate that while corticosterone titer may signal a mother's external and/or internal environment to her developing young, differences in the timing and duration of this endocrine signal produce offspring with specialized phenotypes that exhibit different dispersal strategies.