Female dispersal and reproductive success in wild western lowland gorillas (<Emphasis Type="Italic">Gorilla gorilla gorilla</Emphasis>)

This paper presents data on the dispersal patterns and reproductive success of western lowland gorilla females from a long-term study at Mbeli Bai in the Nouabalé-Ndoki National Park, Republic of Congo. We find that female natal and secondary transfer is common. Female immigration rates are negatively related to group size, and emigration rates are positively related to group size, with the net result that larger groups are losing females and smaller groups are gaining females. Furthermore, females transferring between known groups show a preference for significantly smaller groups. However, there is no effect of group size on female reproductive success. Male protection and male quality are considered important in determining female transfer decisions. The case for infanticide is argued and females exhibit strategies that appear to minimise the probability of infanticide following the death of the silverback. Exclusively single-male groups and group formation through female acquisition by solitary males may bias female transfer to lone silverbacks and small groups. The effects of group size on female dispersal and reproductive success are not wholly consistent with an argument for increased foraging costs, and group size effects are more parsimoniously explained by demographic factors. Male protection from intra-group aggression is the most likely factor underlying grouping patterns across gorilla taxa, but differences in population structure and male reproductive strategies may account for inter-specific variation. We stress the need for intra-specific comparisons and more complete data sets on western lowland gorilla feeding behaviour.Communicated by S. Boinski     

Fitness consequences of dispersal decisions for male mountain gorillas (Gorilla beringei beringei)

The fitness consequences of dispersal decisions are difficult to quantify, especially for long-lived species with complex social systems. To calculate those consequences for male mountain gorillas, from the perspective of both subordinate and dominant males, we used behavioral and demographic data obtained over 30 years from the Virunga Volcano population to develop an agent-based model that simulates the life history events, social structure, and population dynamics of the species. The model included variables for birth rates, mortality rates, dispersal patterns, and reproductive skew. The model predicted an average lifetime reproductive success (LRS) of 3.2 for philopatric males (followers) and 1.6 for emigrants. The benefits of philopatry were most sensitive to opportunities for social queuing and to female transfer preferences, but philopatry remained the best strategy over a wide range of group conditions and hypothetical simulations. The average LRS for dominant males was 4.5 when a subordinate stayed and 4.6 when the subordinate emigrated. The dispersal decision of the subordinate male had little impact on the fitness of the dominant male because it came relatively late in the dominant males reproductive life span, and it changed his group composition only incrementally. The fitness consequences for the dominant male were most sensitive to the degree of reproductive skew. Since subordinates suffer a fitness loss when they leave a group, they should accept whatever reproductive restraint is needed to avoid eviction, and the dominant male does not need to offer concessions for them to stay. The dominant male may offer reproductive concessions for other reasons, such as peace incentives or to confuse paternity, or he may not have complete control of reproduction within his group.Communicated by D. Watts     

Population trend alters the effects of maternal dominance rank on lifetime reproductive success in yellow baboons (<Emphasis Type="Italic">Papio cynocephalus</Emphasis>)

We evaluated the association between dominance rank and lifetime reproductive success of 75 free-ranging female baboons in Mikumi National Park, Tanzania. Data were evaluated over a 22-year period that included a period of troop increase (1975–1987) associated with two troop splits in 1978 and 1979, followed by a precipitous population crash (1987–1996) where the troops successively fused back together in 1989 and 1994. Lifetime reproductive success was significantly greater for high- versus low-ranking females when examined across the entire study period. High-ranking females had a longer reproductive life span (7.4 vs 3.6 years after first birth), reached menarche earlier (4.6 vs 5.2 years), lived longer (12.0 vs 8.8 years), and had more offspring of both sexes (2.25 vs 1.33 for male offspring; 3.25 vs 0.94 for female offspring), with four times the number of offspring of each sex surviving to 4 years of age compared to low ranking females. Greater offspring production was associated with shorter interbirth intervals of dominant versus subordinate females (545 vs 723 days), partly owing to lower miscarriage rates (0.05 vs 0.2) and a shorter duration of lactation (244 vs 330 days). Rank effects were then partitioned by mothers experiencing the majority of their reproductive life prior to, versus during, the population decline. The majority of rank effects on measures of lifetime reproductive success were virtually eliminated for mothers reproducing during the troop decline, indicating that the considerable impacts of social status on lifetime reproductive success can be markedly altered by intrinsically and extrinsically mediated demographic events.Ramon Rhine is deceasedCommunicated by C. Nunn     

The cost of success: reproductive effort in male southern elephant seals (<Emphasis Type="Italic">Mirounga leonina</Emphasis>)

Reproductive effort is a key parameter of life history because it measures the resources allocated to reproduction at the expense of growth and maintenance. Male reproductive effort always had a minor role with respect to female effort both in the development of theories and in field research. Elephant seals are an ideal subject for reproductive effort studies because they fast during the breeding season, splitting the phase of energy acquisition from the phase of energy use for breeding. In this paper, we present results on male reproductive effort (weight loss estimated by photogrammetry) in southern elephant seals (Mirounga leonina), the most dimorphic and polygynous of all mammal species. We show that total reproductive effort increases with age, with no sign of late decrease or senescence. Male reproductive effort in this species depends mostly on behavioral factors, i.e., the success in competition with other males, and the intensity of interaction with females. A large effort results in large gains in both mating success and fertilizations. The large reproductive success that a few males are able to achieve come at a big cost in terms of energy expenditure, but this cost does not seem to affect the likelihood of survival to the following breeding season.     

Cooperative breeding increases reproductive success in the social spider <Emphasis Type="Italic">Stegodyphus dumicola</Emphasis> (Araneae,Eresidae)

Sociality in some birds, mammals, and social insects was suggested to have evolved through the lengthening and extension of parental care behaviors to nondirect descendents. In these systems, group members care for young cooperatively and, thus, increase the reproductive success of the breeders and fitness of the young. Parental care behaviors, such as regurgitation feeding and matriphagy (consumption of the mother), occur in several subsocial and social spiders. However, it is not known whether females in a colony cooperate in caring for the young of other females and whether such cooperative care improves reproductive success. To answer this question, we created experimental colonies of the social spider Stegodyphus dumicola (Araneae, Eresidae), allowing only one female in a group to produce young, simulating reproductive skew occurring in nests in nature. In this paper, we show for the first time that females of S. dumicola cooperate in providing regurgitated food for young of other females and are even eaten by those young. Young raised by a group of females were larger and had greater survival than young raised only by their mother. Thus, fitness benefits from raising broods cooperatively may have favored the evolution of sociality in spiders.     

Helpers in colonial cooperatively breeding sociable weavers <Emphasis Type="Italic">Philetairus socius</Emphasis> contribute to buffer the effects of adverse breeding conditions

Some studies on the effects of helpers in cooperatively breeding vertebrates show a positive effect of helper presence on reproductive output whereas others find no effect. One possibility for this discrepancy is that helpers may have a positive effect when breeding conditions are adverse, while their effect might go unnoticed under good conditions. We investigate this hypothesis on sociable weavers Philetairus socius, a colonial cooperatively breeding passerine that inhabits a semi-arid region where breeding conditions vary markedly. We used multivariate mixed models to analyse the effect of helpers on reproduction under contrasting environmental and social conditions while controlling for parental and colony identity. We found that reproductive success in sociable weavers was primarily influenced by nest predation and rainfall. In addition, colony size was negatively associated with hatching and fledging success and number of young fledged per season. Helpers had a less prominent but significant influence on feeding rates and reproductive outcome. In agreement with expectations, the presence of helpers counteracted some of the negative effects of breeding in periods of low rainfall or in large colonies and was also associated with an increased number of young fledged per season. Our results illustrate that the effect of helpers might be detectable mostly under unfavourable conditions, but can contribute to improve reproductive performance in those situations.     

Rank-related partner choice in the fission–fusion society of the spotted hyena (<Emphasis Type="Italic">Crocuta crocuta</Emphasis>)

When social partners vary in their relative value, individuals should theoretically initiate partnerships with conspecifics of the highest value. Here, we tested this prediction in a wild population of spotted hyenas (Crocuta crocuta). Crocuta live in complex, fission–fusion societies structured by dominance hierarchies in which individuals vary greatly in their value as social companions. Because patterns of association among Crocuta reflect social preferences, we calculated association indices (AIs) to examine how social rank influences intrasexual partner choice among unrelated adults of both sexes. The highest-ranking individuals were generally most gregarious in both sexes. Females associated most often with dominant and adjacent-ranking females. Females joined subgroups based on the presence of particular conspecifics such that subordinates joined focal females at higher rates than did dominants. Dominants benefit from associations with subordinates by enjoying priority of access to resources obtained and defended by multiple group members, but the benefits of these associations to subordinates are unknown. To investigate this, we tested three hypotheses suggesting how subordinates might benefit from rank-related partner choice among unrelated females. We found that subordinates who initiated group formation benefited by gaining social and feeding tolerance from dominants. However, rates at which dominants provided coalitionary support to subordinates did not vary with AIs. Overall, our data resemble those documenting patterns of association among cercopithecine primates. We consider our results in light of optimal reproductive skew theory, Seyfarth’s rank attractiveness model, and biological market theory. Our data are more consistent with the predictions of Seyfarth’s model and of biological market theory than with those of skew theory.     

Sex-specific maternal effect on egg mass,laying order,and sibling competition in the Bengalese finch (<Emphasis Type="Italic">Lonchura striata</Emphasis> var. <Emphasis Type="Italic">domestica</Emphasis>)

Maternal effects, such as investment in eggs, have profound effects on offspring fitness. Mothers are expected to skew their investment depending on the laying order and sex when unequal sibling competition occurs within a brood because of sex-specific vulnerability and age hierarchy caused by asynchronous hatching. The Bengalese finch hatches asynchronously and shows a moderate reversed sexual size dimorphism. However, contrary to commonly accepted assumptions of size-dependent vulnerability, the smaller sex (male) is more vulnerable to developmental stress caused by sibling competition. We investigated whether maternal investment would be biased by the position in laying order and the sex of eggs, and also explored the possible differences in growth patterns depending on sex, laying order, and age hierarchy by observing chicks fostered to experimentally manipulated broods where brood composition was controlled and age hierarchy was more enhanced than in natural breeding conditions. We found that overall patterns of maternal investment favored the disadvantageous sectors of sibling competition, i.e., eggs of later laying order and sons over those of early laying order and daughters. We also examined the effect of laying order on adult body size and sex differences in growth patterns. When reared in the subordinate age hierarchy, females could compensate for the deficit of decreased growth rate by taking longer to mature, whereas males could not. We suggest that this sex-specific growth pattern could be the cause of sex differences in vulnerability to early developmental stress.     

Settlement of the wreckfish (<Emphasis Type="Italic">Polyprion</Emphasis><Emphasis Type="Italic">americanus</Emphasis>)

The wreckfish (Polyprion americanus) is a panoceanic, long-lived, deep-water demersal species, characterized by an extended pelagic juvenile phase associated with floating objects. In the present study morphometric data from 435 specimens collected from Greece (Crete), Italy (Ionian Sea) and the Atlantic coast of France from September 1999 to March 2001 were analyzed to estimate the settlement size of the species and associated changes in morphology. The fishery sample included specimens from both the pelagic and the demersal stage. Length-at-settlement (TL₅₀) and 95% confidence intervals were estimated by fitting a logistic function. Eleven morphometric characters were analyzed, and the existing inflection points, reflecting alterations in body shape, were defined: (1) iteratively, as the transition point, splitting the data set into two groups, for which the reduced major axis functions, between a character and total length, of the successive groups best fit (minimum sum of squares of residuals) the combined data set and (2) where the second derivative of the fitted third-order polynomial functions to morphometric ratios equaled zero. The main size range within which wreckfish settled was 56-65 cm, and the means of the inflection points defined by the two methods were 61 and 64 cm. These results indicate changes in functional morphology associated with settlement. The monitoring of wreckfish juveniles caught in the wild and kept individually in captivity showed changes in growth and food intake at approximately the estimated settlement length and suggested that temperature was the most likely factor triggering settlement.     

Vocal alliances in Chacma baboons (<Emphasis Type="Italic">Papio hamadryas ursinus</Emphasis>)

Theory predicts that females in species with matrilineal dominance hierarchies should use nepotistic support systems to maintain their family’s rank. Female Old World monkeys, however, form alliances against other females at surprisingly low rates. Nonetheless, in many species, females utter threat vocalizations when observing others’ disputes, suggesting that these vocalizations may function as ‘vocal alliances’. We describe a playback experiment testing the efficacy of vocal alliances in free-ranging female baboons. Subjects were played the same female’s threat-grunts under three separate conditions: after being threatened by the signaller’s close relative to mimic kin support, after being threatened by a female maternally unrelated to the signaller to mimic non-kin support, and after a friendly interaction with the signaller’s close relative as a control. Subjects responded more strongly to the playback and avoided the signaller and her matrilineal relatives for a longer period of time in kin support trials than in either non-kin support or no aggression trials. In contrast, there was no difference in subjects’ behaviour between non-kin support and no aggression trials. These results corroborate observational data showing that vocal support occurs at a higher rate than physical support in female baboons, and that kin are more likely to provide vocal support than non-kin. We conclude that vocal support plays a similar role as physical support in the alliances of female baboons.     

Dynamics of pigment degradation by the copepodite stage of <Emphasis Type="Italic">Pseudodiaptomus</Emphasis><Emphasis Type="Italic">euryhalinus</Emphasis> feeding on <Emphasis Type="Italic">Tetraselmis</Emphasis><Emphasis Type="Italic">suecica</Emphasis>

Short-term (3 h) changes in concentration of chlorophylls and their derivatives in stage V Pseudodiaptomus euryhalinus and their fecal material were followed by HPLC during a 24 h experiment. Copepodites were fed with the prasinophyte Tetraselmis suecica. Intact chlorophyll a and b were found in animals and fecal material and had similar dynamics of accumulation over time. The extent of transformation of chlorophyll a and b to colorless compounds was different with chlorophyll a being more extensively degraded. Additionally, several chlorophyll derivatives (pheophytin and pyropheophytin-like pigments) were found. Pyropheophytin a was the most abundant followed by pheophytin b, pheophytin a, and pheophorbide a. Relative amounts of pheopigments were different in copepodites and fecal material, and pheophytin a, pheophorbide a, and pheophytin b concentrations were low and variable. The amount of ingested chlorophyll recovered as chlorophyll a and its derivatives in fecal and copepodite pools was generally low (<5%), with one exception occurring after 9 h, when it accounted for >70%. These data suggest individual pheopigments are produced at different rates and that chemical or enzymatic mechanisms in the gut of copepodites act on the two chlorophylls in different ways.     

Cardiac output during exposure to <Emphasis Type="Italic">Chattonella</Emphasis><Emphasis Type="Italic">marina</Emphasis> and environmental hypoxia in yellowtail (<Emphasis Type="Italic">Seriola</Emphasis><Emphasis Type="Italic">quinqueradiata</Emphasis>)

The experiments were undertaken to measure, for the first time, cardiac output in yellowtail (Seriola quinqueradiata) during exposure to a harmful red tide flagellate (Chattonella marina). The responses were compared with those during exposure to environmental hypoxia to evaluate the significance of the drop of arterial oxygen partial pressure (PaO₂) in the fish-kill mechanisms by C. marina. PaO₂ immediately decreased, whereas heart rate (HR) was maintained until shortly before death during exposure to C. marina. Suffocation developing during the exposure resulted from a decrease in blood oxygen content, but not from lowered blood flow to the tissue. Although exposure to both C. marina and hypoxia immediately decreased PaO₂, arterial oxygen content (CaO₂) and pH (pHa) were significantly lower, whereas HR and cardiac output (&Qdot;) remained significantly higher, for the C. marina-exposed fish than for hypoxia-exposed fish. Although the drop in PaO₂ appears to play a pivotal role in the mechanisms of fish death by C. marina, other physiological response(s) should also be considered.     

Individual specialization in the hunting wasp <Emphasis Type="Italic">Trypoxylon</Emphasis> (<Emphasis Type="Italic">Trypargilum</Emphasis>) <Emphasis Type="Italic">albonigrum</Emphasis> (Hymenoptera,Crabronidae)

Individual-level variation in resource use occurs in a broad array of vertebrate and invertebrate taxa and may have important ecological and evolutionary implications. In this study, we measured the degree of individual-level variation in prey preference of the hunting wasp Trypoxylon albonigrum, which inhabits the Atlantic Forest in southeastern Brazil. This wasp captures several orb-weaving spider genera to provision nests. Individuals consistently specialized on a narrow subset of the prey taxa consumed by the population, indicating the existence of significant individual-level variation in prey preferences. The population niche was broader in the wet season in terms of both prey size and taxa. In the case of prey size, the population niche expansion was achieved via increased individual niche breadths, whereas in the case of prey taxa, individual niches remained relatively constrained, and the population niche expanded via increased interindividual variation. The observed pattern suggests the possibility of functional trade-offs associated with the taxon of the consumed prey. The nature of the trade-offs remains unknown, but they are likely related to learning in searching and/or handling prey. We hypothesize that by specializing on specific prey taxa, individuals increase foraging efficiency, reducing foraging time and ultimately increasing reproductive success.     

Exogenous testosterone increases female aggression in the European starling (<Emphasis Type="Italic">Sturnus vulgaris</Emphasis>)

In the European starling, Sturnus vulgaris, optimal mating systems differ between males and females. Males gain from polygyny, whereas monogamy increases female fitness. The cost of polygyny to females lead to intense female–female competition, and it has previously been shown that the intensity of female aggression during the pre-breeding period can predict the realised mating system. The physiological regulation of such female aggression in starlings is not yet known. This study examines the role of testosterone in mediating aggressive behaviours involved in intra-specific reproductive competition in female starlings. Testosterone levels were experimentally elevated with testosterone implants in females during the pre-laying period. To simulate a situation in which an additional female tried to mate with the focal female’s mate, a caged female was presented close to a nest-site to which the male could attract a secondary female. Testosterone was significantly related to several behaviours involved in female–female interactions. Females with testosterone implants spent significantly more time close to the caged female and produced more song bouts than control females. In contrast, male behaviour was unrelated to the experimental status of the mate. Females mated to males that attracted a secondary female were less aggressive towards the caged female than those that remained monogamously mated. The effect of exogenous testosterone in this study indicates that androgens may mediate social behaviours in female starlings during the breeding season.     

Effects of substrate type on growth and mortality of blue mussels (<Emphasis Type="Italic">Mytilus</Emphasis><Emphasis Type="Italic">edulis</Emphasis>) exposed to the predator <Emphasis Type="Italic">Carcinus</Emphasis><Emphasis Type="Italic">maenas</Emphasis>

Structure and complexity of the substrate are important habitat characteristics for benthic epifauna. The specific growth and mortality rates and inducible defence characters on medium-sized blue mussels (Mytilus edulis L.) exposed to shore crabs (Carcinus maenas L.) were examined on three different substrate types in combined field and laboratory experiments. The experiments showed that complexity of the substrate increased blue mussel survival significantly, through a decrease in predation pressure. However, increased intraspecific competition for food on the complex substrate resulted in significantly lower growth rates of the mussels. Inducible defence characters were also influenced by substrate type. Blue mussels were more affected by predators on the structurally simple substrate, where they developed thicker shells and a larger posterior adductor muscle.     

Viable algae released by the seastar <Emphasis Type="Italic">Dermasterias</Emphasis><Emphasis Type="Italic">imbricata</Emphasis> feeding on the symbiotic sea anemone <Emphasis Type="Italic">Anthopleura elegantissima</Emphasis>

Echinoderms are major predators of anemones in temperate ecosystems. The fate of two algae, zooxanthellae and zoochlorellae, after their host anemone (Anthopleura elegantissima Brandt) was consumed by the leather star Dermasterias imbricata Grube was determined in experiments conducted in July and August 2004. Productivity, photosynthetic pigments, and mitotic index (percent of cells dividing) were used as indicators of algal health; algae released after leather stars consumed their host were compared with algae freshly isolated from anemones. Two types of waste products contained algae: pellets resulting from extraoral digestion, and feces. Zooxanthellae and zoochlorellae isolated from these waste products were photosynthetic, although to different extents. For algae from feces and pellets, light-saturated photosynthetic rates (P _max) were 85 and 13%, respectively, of P _max of freshly isolated zooxanthellae; and were 20 and 46%, respectively, for zoochlorellae. The photosynthetic pigments and mitotic index (percent of dividing cells) were not altered by the feeding activities of the leather star. These results show that algae released by seastar predation on their hosts remain viable, and are hence available for establishing symbioses in A. elegantissima and other potential hosts.     

Antennal responses to floral scents in the butterflies <Emphasis Type="Italic">Inachis io</Emphasis>, <Emphasis Type="Italic">Aglais urticae</Emphasis> (Nymphalidae), and <Emphasis Type="Italic">Gonepteryx rhamni</Emphasis> (Pieridae)

Summary. To better understand the biological role of floral scents for butterflies, electrophysiological responses to floral scents were investigated using combined gas chromatography and electroantennographic detection (GC-EAD). The antennal responses of three butterfly species, Aglais urticae L. (Nymphalidae), Inachis io L. (Nymphalidae), and Gonepteryx rhamni L. (Pieridae) to floral scent compounds from both natural and synthetic mixtures were examined. Floral scents were collected from the butterfly nectar plants Cirsium arvense (L.) (Asteraceae), and Buddleja davidii Franchet cv. (Loganicaeae) with dynamic head-space methods on Tenax-GR and eluted with pentane. These eluates, composed of natural floral scent blends, represent an array of compounds in their natural state. In the GC-EAD analyses eleven compounds were identified from C. arvense with the benzenoid compound phenylacetaldehyde in highest abundance. Seventeen compounds were identified from B. davidii with the irregular terpene oxoisophorone in highest abundance. Thirty-nine synthetic floral scent compounds were mixed in pentane, in equal amounts; about 35 ng were allowed to reach the antennae. The butterflies showed antennal responses to most of the floral scent compounds from both natural and synthetic blends except to the highly volatile monoterpene alkenes. Certain benzenoid compounds such as phenylacetaldehyde, monoterpenes such as linalool, and irregular terpenes such as oxoisophorone, were emitted in relatively large amounts from C. arvense and B. davidii, and elicited the strongest antennal responses. These compounds also elicited strong antennal responses when present in the synthetic scent blends. Thus, the butterflies seem to have many and /or sensitive antennal receptors for these compounds, which points to their biological importance. Moreover, these compounds are exclusively of floral scent origin. For B. davidii, which depends highly on butterflies for pollination, the exclusive floral scent compounds emitted in high abundance could be the result of an adaptive pressure to attract butterflies. Received 2 Septemter 2001; accepted 9 September 2002.     

Foraging responses in the butterflies <Emphasis Type="Italic">Inachis io</Emphasis>, <Emphasis Type="Italic">Aglais urticae</Emphasis> (Nymphalidae), and <Emphasis Type="Italic">Gonepteryx rhamni</Emphasis> (Pieridae) to floral scents

Summary. For butterflies to be efficient foragers, they need to be able to recognize rewarding flowers. Flower signals such as colours and scents assist this recognition process. For plant species to attract and keep butterflies as pollinators, species-specific floral signals are crucial. The aim of this study is to investigate foraging responses to floral scents in three temperate butterfly species, Inachis io L. (Nymphalidae), Aglais urticae L. (Nymphalidae), and Gonepteryx rhamni L. (Pieridae), in behavioural choice bioassays. The butterflies were allowed to choose bet-ween flower models varying in scent and colour (mauve or green). Flowers or vegetative parts from the plants Centaurea scabiosa L. (Asteraceae), Cirsium arvense (L.) (Asteraceae), Knautia arvensis (L.) (Dipsacaceae), Buddleja davidii Franchet (Loganicaeae), Origanum vulgareL. (Lamiaceae), Achillea millefolium L. (Asteraceae), and Philadelphus coronarius L. (Hydrangiaceae) were used as scent sources. All visits to the models — those that included probing and those that did not — were counted, as was the duration of these behaviours. Both flower-naive and flower-experienced (conditioned to sugar-water rewards, the colour mauve, and specific floral scents) butterflies were tested for their preference for floral versus vegetative scents, and to floral scent versus colour. The butterflies were also tested for their ability to switch floral scent preferences in response to rewards. Flower-naive butterflies demonstrated a preference for the floral scent of the butterfly-favourable plants C. arvense and K. arvensis over the floral scent of the non-favourable plants Achillea millefolium (Asteraceae), and Philadelphus coronarius cv. (Hydrangiaceae). Most of the butterflies that were conditioned to floral scents of either C. arvense, K. arvensis, or B. davidii readily switched theirfloral scent preferences to the one most recently associated with reward, thus demonstrating that floral scent constancy is a result from learning. These findings suggest that these butterflies use floral scent as an important cue signal to initially identify and subsequently recognize and distinguish among rewarding plants. Received 2 September 2001; accepted 9 September 2002.