On the social structure of offspring rearing in the burrower bug,<Emphasis Type="Italic"> Sehirus cinctus</Emphasis> (Hemiptera: Cydnidae)

Maternal-offspring interactions are important in a variety of animals. Understanding the evolution of these interactions requires that we also study the broader social context in which they occur. To date, behavioral studies on burrower bugs, Sehirus cinctus, have focused exclusively on interactions between mothers and offspring. Here we ask whether these interactions occur in a social context that extends beyond the family unit of a mother and her own genetic offspring. Such social structure can arise from behaviors that occur before eggs are laid, or from actions of individuals that occur post-hatching. We present field data showing that lay sites of mothers are spatially aggregated on a scale that would lead to behavioral interactions among families. Microsatellite markers suggest neighboring mothers are unrelated. Laboratory experiments do not support the hypothesis that spatial aggregation results from a direct attraction of females to one another. Other laboratory studies reported here indicate that, after hatching, unrelated clutches sometimes join together to form multifamily groups. Experiments reveal that mothers are not necessary for these joining events to occur. In sum, these data suggest that both mothers and offspring play active, but different, roles in generating the social environment in which offspring rearing occurs.Communicated by N. Wedell     

Why are larger convict cichlid (Cichlasoma nigrofasciatum) fry sometimes adopted into broods of smaller fry?

Females of Elasmucha grisea defend their eggs and small nymphs against invertebrate predators. Females sometimes guard their clutches side by side on the same birch leaf. We studied benefits of this joint guarding both in the field and in the laboratory. We found that adjacent females had significantly larger clutches than solitary females. In the laboratory, we studied the effectiveness of joint versus single defence against ant (Formica uralensis) predators. We established female pairs from initially singly guarding females by cutting off pieces of leaves with egg clutches and pasting them beside another female guarding her clutch. In the control group the females with their clutches were similarly cut off but these clutches were placed on another leaf without any female. The birch twigs where females guarded their clutches were placed in cages in close proximity to laboratory ant nests. In the experimental treatment, two females guarded their clutches together and at the same nest there was another birch twig without a female. In the control treatment two twigs with one female on each were placed close to another ant nest. Two females defended their clutches significantly more successfully, losing fewer eggs than did the single females. This primitive form of female sociality in parent bugs resembles colonial nesting in birds, where communal defence is also important. However, to our knowledge this is the first experiment where the benefit of joint guarding has been tested directly by manipulating the size of the breeding group rather than by measuring the risk of predation in groups of different size.     

Factors influencing length of maternal care in brown bears (<Emphasis Type="Italic">Ursus arctos</Emphasis>) and its effect on offspring

Length of maternal care, i.e. the interval between successfully raised litters, is the most important factor explaining the variation in reproductive rate among brown-bear (Ursus arctos) populations. In this paper, we examine the variation in length of maternal care in radio-marked brown bears and its effect on their offspring in northern Sweden. Young stayed with their mothers for 1.4–1.5 or 2.4–2.5 (in one case 3.5) years and were weaned with body masses varying from 17 to 69 kg. The probability of yearling litters staying with their mother for a 2nd year increased with decreasing yearling body mass, and was higher for litters with two offspring than for litters with one or three to four offspring. Staying with their mothers for a 2nd year had a positive effect on mass gain in yearlings and this effect was more pronounced in litters with two than three to four offspring. Body mass of 2-year-olds was not related to age of weaning, suggesting that keeping offspring for an additional year mainly compensated for low yearling body mass. If large offspring body mass positively affects later offspring survival and reproduction, mothers may be able to optimize the length of maternal care according to the litter size and the size of their yearlings.Communicated by F. Trillmich     

Prelaying maternal condition modifies the association between egg mass and T cell-mediated immunity in kestrels

Theoretical models of parental care predict differential allocation in relation to the expected fitness of the offspring. As birds must allocate all the resources required for embryonic development in self-contained packages, maternal egg allocation takes a central role in avian life histories. It is predicted that mothers can influence offspring performance by varying the size or composition of their eggs. In this context, T cell-mediated immune response has been revealed as a valuable tool to evaluate fledgling fitness. However, little is known about its relationships to egg mass. I describe a negative association between egg mass and cellular immune response in Eurasian kestrel nestlings (Falco tinnunculus) throughout 3 years. To explore how environmental conditions could mediate this relationship, I experimentally supplied food to females before egg formation in the third study year. I found that this relationship turned to positive in food-supplied females without any effect on egg size. This result shows that environmental conditions before egg formation mediate the relationship between egg mass and future immune response of the nestlings. I suggest that females may be constrained in improving the future immune response of their offspring due to resource allocation trade-offs determined by their pre-laying condition.     

Food abundance and parental care in yellow warblers (Dendroica petechia)

Emlen and Oring (1977) suggested that monogamy in birds is maintained because of the need for strict biparental care. A corollary of their suggestion is that paternal care should decrease under conditions of high food abundance. An alternative is that paternal care would increase if males take advantage of the higher food abundance by trying to reduce the length of the nestling feeding period. We tested these two ideas using yellow warblers (Dendroica petechia) by providing some pairs with supplemental food, thereby reducing the importance of biparental care. However, the extra food did not decrease paternal effort, nor did it increase it (Fig. 2). Early in the nestling period experimental females brooded more but visited their nestlings less than did control females, but later, when brooding times decreased, experimental females fed their nestlings more than did control females (Fig. 3). There were no significant differences in nestling survival (Fig. 5), but nestlings in the control treatment were larger and heavier up to 6 days old (Fig. 6). The main effect of supplemental food was on maternal, not paternal behaviour. Models of biparental care assume interdependence between the parental effort of both parents. In this species, however, males and females provide for their brood independently from each other.     

Tactics, effectiveness and avoidance of mate guarding in the blue-footed booby (Sula nebouxii)

We examined the dynamics and avoidance of mate guarding, by males and females, in the blue-footed booby, in which the two social mates are usually simultaneously present on the territory but each of them is unmonitored by the other for one-quarter of its time. Both sexes were promiscuous and liable to switch mates. Cuckolded individuals did not increase their overall presence on the territory, but in response to the extra-pair (EP) courtships of their mates, both sexes doubled their rate of intra-pair (IP) courtship and sometimes showed aggression. The male or female's presence depressed the social mate's EP activity, but intra-pair courtship had no such effect, tending even to propitiate that EP activity. Similarly, when females responded to their social mates' EP courtship with approach or aggression, disruption of EP activity was short-lived. Promiscuous females modified their diurnal pattern of attendance, as if attempting to sidestep monitoring by their mates, but cuckolded males matched the modification. Both sexes tended to perform their EP activities at a distance when their mates were present, possibly to evade monitoring or disruption by their mates. Male and female boobies cannot monitor their mates continuously, they do little to facultatively adjust their presence on territory to the risk of infidelity, and their immediate responses to overt infidelity have only the briefest impact; but the information they acquire while monitoring their mates may be critical to constraining their mates' infidelity and also to calibrating their own reproductive investment.     

Reproductive effort in relation to maternal social rank in reindeer (<Emphasis Type="Italic">Rangifer tarandus</Emphasis>)

In polygynous mammals, high-quality females may increase their fitness by providing superior care to their offspring. Based on the agonistic interactions of female reindeer in an experimental herd during two consecutive years (1997 and 1998), we tested whether maternal social rank influenced: (1) winter body-mass change of females, (2) preparturition reproductive effort (measured as fecundity, the birth mass and the birth date of their calves), (3) preweaning maternal effort (measured as calves preweaning mortality, early preweaning and late preweaning growth rate and September body mass of calves), and (4) postweaning maternal effort (measured as calves body-mass change during their first winter). In the models, we included September females body mass as a covariate to separate the effects of maternal rank and body mass. We also tested whether the effect of social rank on maternal efforts was dependent on offspring sex. High-ranked females gained body mass whereas low-ranked females lost weight during the winter. Fecundity was higher and date of birth was earlier in high-ranked females than in subordinates, whereas no effect of females rank on birth mass of calves was found. Early preweaning growth rate and September body mass of calves increased with increasing females social rank, whereas late preweaning daily growth rate of calves was not influenced by females rank. Calves preweaning mortality was only influenced by year, which also explained most of the variance in the winter body-mass change of calves. The effects of females rank on the reproductive-efforts parameters studied were not specific to offspring sex. These findings suggest that females rank influences reproductive effort during the preparturition, as well as the preweaning, period, the effect being sex independent.Communicated by R. Gibson     

Benefits and costs of earwig (Forficula auricularia) family life

The evolution of parental care and family group formation critically depends on offspring survival benefits and parental fecundity costs of care under given ecological conditions. Investigations of the functional significance of care in insect species that exhibit facultative parental care have been relatively rare but may be of particular interest for better understanding of benefit and cost schedules at an early evolutionary stage. In this study, aspects of benefits and costs of care were addressed in the sub-social European earwig (Forficula auricularia; Dermaptera: Forficulidae) by manipulating the presence of tending mothers and brood size in a fully crossed experimental design. Larvae growing in broods tended by their mother or of reduced size showed a higher survival probability than larvae growing in untended or large broods, as predicted if maternal care is beneficial and shaped by a trade-off between number and quality of offspring. Analysis of patterns of food consumption and developmental time further suggested that the benefit of maternal attendance is mediated by the maternal provisioning of food, while the quality–quantity trade-off seemed to be driven by sibling rivalry. Further, tending mothers delayed the production of a second clutch, indicating a potential cost of care in terms of lifetime fecundity. This study experimentally shows benefits and potential costs of maternal care and family group formation in the European earwig. More detailed behavioural experiments will be required to fully understand how behavioural interactions among family members mediate these reproductive outcomes.     

Male sexual attractiveness affects the investment of maternal resources into the eggs in peafowl (<Emphasis Type="Italic">Pavo cristatus</Emphasis>)

According to the differential investment hypothesis, females paired with attractive mates are expected to invest more in the current reproduction relative to females paired with unattractive males. We experimentally tested this hypothesis in the peafowl (Pavo cristatus) by providing females with males that differed in sexual attractiveness. In agreement with the differential allocation hypothesis, females paired with more ornamented males laid larger eggs, and deposited higher amounts of testosterone into the egg yolk, independently of the sex of the embryo. These results show that the association between paternal phenotype and offspring quality could arise via a differential maternal investment. They also suggest that, if ornamented males do transmit good genes to the progeny, the maternal differential investment can amplify the effect of such good genes on the offspring fitness.     

Ecological and hormonal correlates of antipredator behavior in adult Belding’s ground squirrels (Spermophilus beldingi)

Predator–prey relationships provide an excellent opportunity to study coevolved adaptations. Decades of theoretical and empirical research have illuminated the various behavioral adaptations exhibited by prey animals to avoid detection and capture, and recent work has begun to characterize physiological adaptations, such as immune reactions, metabolic changes, and hormonal responses to predators or their cues. A 2-year study quantified the activity budgets and antipredator responses of adult Belding’s ground squirrels (Spermophilus beldingi) living in three different California habitats and likely experiencing different predation pressures. At one of these sites, which is visually closed and predators and escape burrows are difficult to see, animals responding to alarm calls remain alert longer and show more exaggerated responses than adults living in two populations that likely experience less intense predation pressure. They also spend more time alert and less time foraging than adults at the other two sites. A 4-year study using noninvasive fecal sampling of cortisol metabolites revealed that S. beldingi living in the closed site also have lower corticoid levels than adults at the other two sites. The lower corticoids likely reflect that predation risk at this closed site is predictable, and might allow animals to mount large acute cortisol responses, facilitating escape from predators and enhanced vigilance while also promoting glucose storage for the approaching hibernation. Collectively, these data demonstrate that local environments and perceived predation risk influence not only foraging, vigilance, and antipredator behaviors, but adrenal functioning as well, which may be especially important for obligate hibernators that face competing demands on glucose storage and mobilization.     

Compensating for the costs of polygyny in hen harriers Circus cyaneus

In polygynous species, the adults are faced with a dilemma during chick rearing. Males must decide how to distribute food between their females and food allocation patterns are often highly unequal. In turn, the females that receive less food from males have to decide how much time to invest in additional hunting. If they spend more time hunting, then they leave their young exposed to weather and predators. However, if they stay at the nest, they increase the risk of their chicks starving. One way that birds may compensate for reduced provisioning is by increasing the size of prey caught. We tested this hypothesis by comparing prey deliveries to nests of hen harriers, Circus cyaneus, with females of different breeding status. As expected, male harriers delivered less food items to the nests of polygynous females, and especially their secondary, or β females. However, both sexes were able to compensate by delivering larger items and there was no difference in the overall mass of food delivered to nests. Moreover, females spent a similar amount of time at the nest, irrespective of status, and there were no overall differences in breeding success. Our results show that polygynous female harriers can compensate for the costs of polygyny, but we suggest that their ability to do so will vary according to the abundance of both large prey and predators.     

Territorial vocal behavior in hybrid smooth froglets, Geocrinia laevis complex (Anura: Myobatrachidae)

Males of the parapatically distributed myobatrachid frogs Geocrinia laevis and G. victoriana have highly divergent advertisement calls. Furthermore, the two species differ strongly in the complexity of their vocal repertoires, with males of G. victoriana possessing, and those of G. laevis lacking, a distinct territorial vocalization (encounter call). We investigated the territorial vocal behavior of males in a persistent natural hybrid population. Most hybrid males possessed encounter calls functionally equivalent to those of G. victoriana, that were produced following exposure to playback of recorded advertisement calls presented at >110 dB peak sound pressure level. The territorial acoustic responses were not associated with an index of hybridity derived from the structure of the advertisement call, suggesting genetic and functional decoupling of the two components of the vocal repertoire; i.e., advertisement calls and encounter calls. This decoupling may be the result of sexual selection favouring those hybrids with pronounced territorial behavior and the associated vocalization, regardless of the structure of their advertisement calls.Communicated by A. Mathis     

Infant care,infanticide, and female reproductive strategies in polygynous groups of common marmosets (Callithrix jacchus)

Reproduction in marmoset and tamarin groups is typically restricted to a single dominant female, but it is unclear why subordinate females tolerate delayed reproduction. The presence of two breeding females in free-ranging groups of common marmosets (Callithrix jacchus) presented a unique opportunity to examine differences in the reproductive strategies of dominant and subordinate females. Three groups were monitored for 12–18 months at a forest reserve in northeastern Brazil. Data on infant care were collected during two consecutive all-day follows every 10 days until the infants were 2 months old. Carrying patterns for infants born to dominant females were similar to those observed in groups containing a single breeding female. All group members over 5 months of age participated in infant care, and dominant females allowed some group members to carry their infants from the 1 st day of life. In contrast, subordinate females were protective of their offspring and were their sole caretakers for at least a week following birth. One infant born to a subordinate was killed in an attack involving the dominant female. Overall, dominant females gave birth to more infants and had higher infant survival than did subordinate breeding females. subordinate females were successful in rearing young only when the timing of births was such that they did not overlap with the dependency period of infants born to the dominant female. These patterns suggest that subordinate females may face resource competition, especially over access to helpers. In times of severe competition, subordinate females that delay reproduction may be avoiding a wasted reproductive effort.     

Male care of infants in a siamang (Symphalangus syndactylus) population including socially monogamous and polyandrous groups

While male parental care is uncommon in mammals, siamang (Symphalangus syndactylus) males provide care for infants in the form of infant carrying. I collected behavioral data from a cohort of five wild siamang infants from early infancy until age 15–24 months to identify factors affecting male care and to assess the consequences of male care for males, females, and infants in a population including socially monogamous groups and polyandrous groups. There was substantial variation in male caring behavior. All males in polyandrous groups provided care for infants, but males in socially monogamous groups provided substantially more care than males in polyandrous groups, even when the combined effort of all males in a group was considered. These results suggest that polyandry in siamangs is unlikely to be promoted by the need for “helpers.” Infants receiving more care from males did not receive more care overall because females compensated for increases in male care by reducing their own caring effort. There was no significant relationship between indicators of male–female social bond strength and male time spent carrying infants, and the onset of male care was not associated with a change in copulation rates. Females providing more care for infants had significantly longer interbirth intervals. Male care may reduce the energetic costs of reproduction for females, permitting higher female reproductive rates.     

Changes in singing behavior of male black-capped chickadees (Parus atricapillus) following mate removal

We removed the mates of ten male black-capped chickadees (Pares atricapillus) during the nest-building period to determine the effect of female presence on dawn singing. During the first dawn chorus following mate removal, males sang significantly longer, increased movement within their territory, and increased the percentage of their territory covered while singing. After the female was returned, these parameters returned to the pre-removal values. Males did not alter the frequency range or modal frequency of their songs when the mate was removed, nor did they change the degree of frequency shifting in the fee-bee song. We conclude that dawn singing in the black-capped chickadee acts, in part, as an intersexual signal, and that the behavior of frequency shifting in the song may be directed more toward rival males than females. Correspondence to: K. Otter     

Benthic macrophyte communities in the Ghar El Melh lagoon (North Tunisia)

This work deals with the biodiversity and distribution of benthic macrophytes in the Ghar El Melh lagoon, a Mediterranean coastal lagoon located in the North of Tunisia. An inventory was made of the benthic flora and submerged macrophyte communities were mapped during two successive campaigns (the summer of 1999 and the winter of 2000). The following 24 macrophyte species were identified: seven red algae, two brown algae, 11 green algae, and four marine angiosperms. The results were compared with available data from the literature. Ruppia cirrhosa is the most dominant species. It is found in all lagoon parts, except in the west sector.Ruppia beds are usually associated withCladophora forming heterogeneous communities. During summerRuppia cirrhosa shows a large distribution, covering an area of ca. 21.4 km², with dense, extensive beds covering 80–100%. In winter, severalCladophora species have a very large distribution as well, covering nearly an area of 28.5 km² with an average cover of 46%. The green algaeCaulerpa prolifera is confined to the eastern part of the lagoon which is mainly affected by seawater. In comparison with previous situations, many transformations were observed in biodiversity and spatial distribution of the dominant communities. Thus,Cymodocea nodosa andZostera beds, which dominated in the 1970s, were replaced byZostera andCaulerpa prolifera in the 1980s and are currently succeeded byRuppia cirrhosa andCladophora. Restoration of the Ghar El Melh lagoon will enable an increase in the exchange with the open sea and the circulation of water, in particular in the confined zones. This should considerably improve the water quality and would positively influence the phytobenthic communities.     

Queen dimorphism and reproductive strategies in the fire ant Solenopsis geminata (Hymenoptera: Formicidae)

Alate trapping studies of a monogyne population of the fire ant Solenopsis geminata indicate that two sizes of gynes are produced. Macrogynes, which participate in late spring and summer mating flights, are larger, fattier, and more than twice as heavy as microgynes, which participate in fall mating flights. Three patterns of gyne production were observed in 51 colonies studied: 35 produced macrogynes only, 9 produced microgynes only, and 7 produced both morphs, contributing to both summer and fall mating flights. Behavioral evidence and rearing studies suggest that macrogynes found new colonies independently, whereas microgynes achieve colony queen status by infiltrating or being adopted by established colonies. Of the total number of female alates collected from the trapped colonies, 56% were microgynes. However, because of their smaller size and lower fat content, microgynes made up only one-third of the caloric investment in female alates. By measuring the thorax lengths of queens from mature colonies, we determined that at least 56% were macrogynes and 35% or more were microgynes. These results indicate that as a reproductive strategy, colony investment in microgyne production may have at least as high a payoff as investment in macrogyne production.This is publication #24 of the Fire Ant Research Team     

Characterization of the benthic vegetation in the Farwà Lagoon (Libya)

This study forms part of the Action Plan for the Conservation of Marine Vegetation in the Mediterranean SEa (United Nation Environmental Program). It was carried out in June 2000 in the Farwà Lagoon, Libya. The mapping of the main benthic vegetation was achieved by compiling the field observations (transect method), and remote sensing of SPOT satellite images. The phytobenthos in the Farwà lagoon covers an area of 1820 ha (65%). Three benthic macrophyte species dominate, namely the marine phanerogamsCymodocea nodosa andPosidonia oceanica, and the algaCaulerpa prolifera. DeadPosidonia oceanica leaves (litter) form veritable mounds in the vicinity of the openings leading to open sea. These leaves, which come from the coastal sea, are brought into the lagoon by currents and tides; their decomposition will lead to high oxygen consumption and the release of hydrogen sulphide. The phenological data ofPosidonia oceanica shoots sampled in the lagoon are similar to those from other stations in the Mediterranean. Conversely, the lepidochronological parameters of shoots sampled in the central part of the lagoon exhibit values that are substantially higher than those generally recorded in the Mediterranean. The mean number of leaves produced annually is 9.9 (mean value for the Mediterranean: 7.5) and the rhizome growth rate is of 35.7 mm.yr⁻¹ (mean value for the Mediterranean: 7.5 mm.yr⁻¹). This hypersaline environment would seem to provide optimum growth conditions for the speciesPosidonia oceanica.     

High paternal investment in unrelated young: extra-pair paternity and male parental care in house martins

The response of males to reduced paternity has important consequences for the evolution and maintenance of a mixed reproductive strategy. Paternity is predicted to affect directly the level of male parental care in some cases but not in others. The response of males to reduced paternity will be influenced by their ability to assess their paternity, the predictability of cuckoldry and the costs and benefits of parental care. Although male house martins (Delichon urbica) provide among the highest levels of male parental care known in passerines (incubation, brooding and feeding nestlings), there was no evidence that cuckolded males substantially reduced their level of parental care, and, as a result, all young fledged successfully. Thus, extra-pair fertilizations enhanced the reproductive success of some males because they were able to parasitize the parental care of cuckolded males. We discuss several conditions which may favor extensive male parental care even when the male's paternity is very low.     

The effects of larval experience with a complex plant latex on subsequent feeding and oviposition by the cabbage looper moth: Trichoplusia ni (Lepidoptera: Noctuidae)

Summary. Leaf disc choice and oviposition bioassays were used to examine the effects of larval experience with a Hoodia gordonii latex on subsequent behaviors. The latex deterred feeding and oviposition by “naïve” cabbage looper (Trichoplusia ni, Noctuidae) larvae and moths with no previous exposure to the material. “Experienced” insects, reared on a diet with the H. gordonii latex (1000 ppm), exhibited lesser feeding deterrence relative to naïve insects. Experienced female moths actually preferred to lay eggs on treated rather than control leaves. There was no observed transfer of behavioral preferences from experienced parents to their offspring. Our results suggest that moths may be acquiring oviposition preferences from larval feeding experience as described by Hopkins’ host selection principal (HHSP) or through chemical legacy.