Sperm utilization pattern in the honeybee (<Emphasis Type="Italic">Apis mellifera</Emphasis>)

Queen honeybees (Apis mellifera) mate with a large number of drones on their nuptial flights. Not all drones contribute equally to the queens offspring and the queens utilization pattern of spermatozoa from different drones has an important impact on the genetic composition of the colony. Here we study the consequences of sperm use for the fitness of the queens mates with microsatellite DNA-fingerprinting. Eight queens were instrumentally inseminated with semen of six or seven drones. Each drone contributed either 0.5 µl or 1.0 µl semen, respectively, and we analyzed both the impact of the insemination sequence and the amount of semen on the sperm utilization. Our data show no significant effect of the insemination sequence but a strong impact of the semen volume of a drone on the frequency of his worker offspring in the colony. This effect was not linear and the patriline frequencies of the drones contributing larger semen volumes are disproportionately enhanced. If these observations are also valid for natural matings, drone honeybees should maximize the number of sperm but not apply specific mating tactics to be first or last male in a mating sequence.Communicated by R. PageAn erratum to this article can be found at     

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     

Mate guarding,copulation strategies and paternity in the sex-role reversed,socially polyandrous red-necked phalarope<Emphasis Type="Italic"> Phalaropus lobatus</Emphasis>

In a recent review, Westneat and Stewart (2003) compiled evidence that extra-pair paternity results from a three-player interaction in which sexual conflict is a potent force. Sequentially polyandrous species of birds appear to fit this idea well. Earlier breeding males may attempt to use sperm storage by females to obtain paternity in their mates subsequent clutches. Later-breeding males may consequently attempt to avoid sperm competition by preferring to pair with previously unmated females. Females may bias events one way or the other. We examined the applicability of these hypotheses by studying mating behavior and paternity in red-necked phalaropes (Phalaropus lobatus), a sex-role reversed, socially polyandrous shorebird. Male red-necked phalaropes guarded mates more strongly than other shorebirds. Males increased within-pair copulation attempts during their mates fertile period, and maintained or further increased attempts towards the end of laying, suggesting an attempt to fertilize the females next clutch; these attempts were usually thwarted by the female. Paired males sought extra-pair copulations with females about to re-enter the breeding pool. Multilocus DNA fingerprinting showed that 6% of clutches (4/63) each contained one chick sired by a male other than the incubator, producing a population rate of these events of 1.7% (n=226 chicks). Male mates had full paternity in all first clutches (n=25) and 15 of 16 monogamous replacement clutches. In contrast, 3 of 6 clutches of second males contained extra-pair young likely fathered by the females previous mate. Previously mated female phalaropes may employ counter-strategies that prevent later mating males from discriminating against them. The stability of this polyandrous system, in which males provide all parental care, ultimately may depend on females providing males with eggs containing primarily genes of the incubating male, and not a previous mate.Communicated by M. Webster     

Spotted bowerbirds<Emphasis Type="Italic"> Chlamydera maculata</Emphasis> do not prefer rare or costly bower decorations

Females target elaborate secondary sexual traits to acquire either direct benefits, represented or provided by the trait, or indirect benefits usually in the form of genetic components. The link between a males trait and his potential genetic contribution is often unclear. Bowers are extreme examples of secondary sexual traits, built and decorated by males and targeted by females. However, why females should base their mate choice on such structures is unclear. One of the simplest suggestions is that bowers are sites where males exhibit collections of objects that are inherently highly costly, indicating male quality to the choosy female. Such costs can be imposed in two ways, acquisition from the local area, and maintenance in the face of degradation. We studied bowers of spotted bowerbirds Chlamydera maculata at three sites in central Queensland and tested whether components of the bowers that best predict mating success were some of the most costly in the local environment by measuring their abundance, physical dimensions and rate of decay. We also tested whether rare objects per se found on the bowers, were relatively good predictors of mating success. Finally, we experimentally tested whether males preferentially chose rare, and hence costly, objects to use as decorations. We found that objects used on bowers were not unusually costly. Specific objects identified as good predictors of mating success were also some of the more common objects found in the area. They were not the largest objects used on bowers and decayed slowly relative to other objects available for use as decorations. Rare objects in general were not good predictors of mating success and males did not prefer to use rare objects when offered choices. In conclusion, we found no support for the theory that by simply inspecting numbers of decorations and assessing their costs of acquisition and maintenance females can easily obtain information regarding the owners quality. This is due to the fact that decorations used by spotted bowerbirds are not inherently costly.Communicated by J. Graves     

False feedings at the nests of carrion crows<Emphasis Type="Italic"> Corvus corone corone</Emphasis>

False feedings, when individuals visit the nest but refrain from feeding the chicks, occur in some cooperative species and have been interpreted in the white-winged chough (Corcorax melanorhamphos) as active deception by helpers towards the rest of the group. In a cooperatively breeding population of carrion crows (Corvus corone corone) 81.5% of the individuals that provided nestling care showed various kinds of false feedings: arriving at the nest with no food, consuming part or all the food brought to the nest, or taking back from a chicks gape the food that had just been delivered. False feedings occurred on average during 16.3% of nest visits, with some individuals performing them at very high rates (up to 64% of nest visits). False feedings occurred at similar rates in unassisted pairs and groups with helpers, and breeding females showed false feeding at significantly higher rates than other group members. Furthermore, individuals showed false feedings regardless of whether they were alone on the nest or in the presence of other group members, and false feedings did not provoke aggression by the rest of the group. False feedings are not likely to represent deceptive help in the carrion crow. We suggest that crows evaluate the chicks condition during nest visits and that false feedings occur as result of a trade-off between their own hunger and the chicks needs.Communicated by W.A. Searcy     

Pheromones inBlattella germanica andBlaberus craniifer (Blaberoidea): Glandular source,morphology and analyses of pheromonally released behaviours

Females ofBlattella germanica andBlaberus craniifer produce a volatile sex pheromone attractive at a distance for conspecific males. During the emission of the pheromone, females ofB. craniifer adopt a typical calling posture; we never observe such a stance inB. germanica females. For both species, the glandular structures responsible for pheromonal production are located on the female pygidium. InB. germanica, the thickness of the glandular epithelium is clearly correlated with the attractiveness of females, being maximum for 13 day old females. The functions of the pygidial glands are discussed in both species.     

Earlier sex change in infected individuals of the protogynous reef fish<Emphasis Type="Italic"> Thalassoma bifasciatum</Emphasis>

Sex allocation theory for sequential hermaphrodites predicts the size at which an individual should change sex, given the different relationships between individual size and reproductive success in the two sexes. We studied a host-parasite system where the myxozoan Kudoa ovivora infects the ovaries of the reef fish Thalassoma bifasciatum, a protogynous sequential hermaphrodite. The parasite sporulates in the hosts eggs and renders them infertile. It is thus expected to reduce the females reproductive success, and could thereby influence host sex change. We present data from marked fish we observed in the field over 4 months. The data suggest that females infected with Kudoa ovivora have a lower reproductive success, change sex earlier and at a smaller size than uninfected females. These results are in agreement with predictions from sex allocation theory, and provide the first example of a possible parasitic influence on the sex allocation of its host.Communicated by P. Heeb     

Aluminium and fluoride concentrations of three tea varieties growing at Lantau Island,Hong Kong

The present project aims to investigate aluminium (Al) and fluoride (F) contents in teas (Camellia sinensis (L.) O. Kuntze). Three different commercial tea varieties: Assam variety and two China sub-varieties, a large leafed variety and small leafed variety, were collected in two tea gardens of Lantau Island tea plantation of Hong Kong. In general, high concentrations of Al and F were accumulated in the mature leaves (15.3 and of 2.07 g kg^–1 respectively). Among the three varieties, the small leafed variety exhibited the highest Al and F contents followed by the large leafed variety whereas the Assam variety had the lowest Al and F concentrations in its tea bushes. Tea products from a plantation were also analysed and it was noted that black tea had higher Al and F concentrations than green tea. The amount of Al and F released into tea liquor was also tested and the results showed that higher concentrations of Al and F were released into tea liquor under repeated infusion method than continuous infusion method.     

Courtship and reproduction in<Emphasis Type="Italic"> Carybdea sivickisi</Emphasis> (Cnidaria: Cubozoa)

Courtship and fertilization events in cubozoans have received little attention from biologists, and much of what we know about these processes is based on conjecture or scant anecdotal evidence. I set out to describe these processes in the cubozoan Carybdea sivickisi by observing mature medusae in vitro. Mature adults engage in courtship during which spermatophores are transferred from the male to the female, who then inserts the gametes into her manubrium. Females accepted multiple spermatophores from multiple males, and only produced one embryo strand. This study also provides evidence that the presence of conspicuous velar spots on the females bell margin is a signal of sexual maturity, and that sexual maturity was not reached in either sex until individuals had a bell diameter of at least 5 mm.Communicated by R.J. Thompson, St. Johns     

Arrived in the light: diel and seasonal activity patterns in wild Verreaux’s sifakas (<Emphasis Type="Italic">Propithecus v. verreauxi</Emphasis>; Primates: Indriidae)

Madagascar is characterised by pronounced annual climatic and ecological seasonality and harbours a radiation of closely related sympatric primates (Lemuriformes) that exhibit diurnal, nocturnal and cathemeral activity patterns. We collected continuous activity data over a complete annual cycle from wild diurnal Verreauxs sifakas (Propithecus v. verreauxi) to contribute detailed and comparative data: (1) to characterise their diel and seasonal activity rhythms, (2) to identify factors shaping variation in activity rhythms, and (3) to help reconstruct the evolutionary transition from nocturnal to diurnal activity. We fitted eight adult sifakas from seven different groups living in Kirindy forest with an accelerometer/data logger device and recorded their activity in 5-min bins for a total of 12 months. We characterise P. verreauxi as a strictly diurnal species with a pronounced bimodal activity pattern that briefly changed to a more unimodal pattern during their annual mating season (January to March). We documented significant annual variation in total daily activity, activity time, and activity level, as well as in most parameters characterising their bimodal activity pattern. Despite a significant positive correlation of the animals activity time with day length, pronounced annual variation in the phase position of onset and end of activity in relation to sunrise and sunset times could also be discerned. Minor enhancing effects of moonlight on nocturnal activity were only found for the first 3 h of the inactivity period. Bimodality of the activity pattern and the additional reduction of activity time during the cold and dry winter months associated with reduced food availability can be interpreted as flexible behavioural adaptations to reduce energy expenditure. We therefore propose that energetic and thermoregulatory benefits are important factors shaping these primates activity pattern.Communicated by C. Nunn     

Does the presence of non-breeders enhance the fitness of breeders? An experimental analysis in the clown anemonefish<Emphasis Type="Italic"> Amphiprion percula</Emphasis>

The stability of animal societies depends on individuals decisions about whether to tolerate or evict others and about whether to stay or leave. These decisions, in turn, depend on individuals costs and benefits of living in the group. The clown anemonefish, Amphiprion percula, lives in groups composed of a breeding pair and zero to four non-breeders. To determine why breeders accept the presence of non-breeders in this species I investigated the effect of non-breeders on multiple components of the breeders fitness. Non-breeders did not assist breeders in any obvious way. Experimental removal of non-breeders had no significant effect on the survival, growth, or reproductive success of breeders. Experimental removal of one of the breeding pair showed that non-breeders had little effect on the time taken for a widowed breeder to recommence breeding. The results indicate that the presence of non-breeders neither enhances, nor reduces, the fitness of breeders in A. percula. I suggest that non-breeders might modulate their effect on the fitness of breeders, either by reducing the costs they inflict or by increasing the benefits they provide, such that it just pays breeders to tolerate, rather than to evict, them. This study illustrates that animal societies can be stable even when some individuals gain nothing from the association.Communicated by M. Abrahams     

Sexual dimorphisms in radula of<Emphasis Type="Italic"> Conomurex persicus</Emphasis> (Gastropoda: Strombidae) in the Mediterranean Sea

Dentition characters of Conomurex persicus found along the Mediterranean coasts of Turkey were analyzed. For the analyses of radula, 289 individuals (34 adult males, 140 adult females and the remainder juveniles) were examined. Male and female individuals of the species had different numbers of radular cusps. The radula of the taenioglossate type was made up of two marginal teeth, one lateral and one central (rhachidian) tooth, each being delicate and with different numbers of cusps. The central and lateral teeth had seven cusps and five cusps, respectively, in both males and females. Outer and inner marginal teeth of males had two and one cusps, while those of females had five and six cusps, respectively. The females radula is yellow-whitish. The males radula is reddish dark brown.Communicated by O. Kinne, Oldendorf/Luhe     

Patterns of participation and free riding in territorial conflicts among ringtailed lemurs (<Emphasis Type="Italic">Lemur catta</Emphasis>)

Cooperation in animal social groups may be limited by the threat of free riding, the potential for individuals to reap the benefits of other individuals actions without paying their share of the costs. Here we investigate the factors that influence individual contributions to group-level benefits by studying individual participation in territorial defense among female ringtailed lemurs (Lemur catta). To control for potentially confounding factors, particularly group size, we studied two semi-free-ranging groups at the Duke University Primate Center. First, we used a combination of experimental and observational methods to investigate the costs and benefits of territorial defense for individual lemurs. We found three indications of costs: physical contact occurred during inter-group encounters, participation in territorial defense was negatively correlated with ambient temperature, and rates of self-directed behaviors increased during encounters. Benefits were more difficult to quantify, but observational and experimental tests suggested that individuals shared the gains of territorial defense by foraging in defended territories. Thus, during experiments in which one of the groups was prevented from defending its territory, the free-ranging group made more frequent incursions into the other groups territory. Second, we examined variation in participation in territorial defense. Individuals varied significantly in their rates of aggression and genital marking during inter-group encounters. The extensive variation documented among individuals was partially accounted for by dominance rank, kinship and patterns of parental care. However, we found no evidence to suggest that participation was enforced through punishment (policing) or exchange of benefits involving grooming. In conclusion, this study provides further insights into cooperative behavior in mammalian social groups by revealing how the costs and benefits of territoriality influence patterns of individual participation in the context of shared (collective) goods.Communicated by P. Kappeler     

Parasitism can influence the intertidal zonation of non-host organisms

We investigated habitat utilization in Blainvilles beaked whale, Mesoplodon densirostris, in the northern Bahamas and, as such, this is the first analysis of fine-scale habitat utilization of any member of the genus Mesoplodon. We divided the area into 500×500 m grid squares and each square was assigned a classification for presence or absence of Blainvilles beaked whales, along with details of water depth, seabed gradient and seabed aspect. All squares where Blainvilles beaked whales occurred had gradients from 68 to 296 m/km and depths from 136 to 1,319 m and most faced northeast compared with 0–526 m/km, 10–3,000 m and all aspects for the whole study area. Generalized additive models and classification trees indicated that, in order of importance, aspect, gradient and depth were related to occurrence of Blainvilles beaked whales within the study area. We hypothesize that the relationships between habitat utilization and these topographic variables relates to the effects of interactions between a deepwater current and the seabed topography on preferred prey. Locally, prey animals may be concentrated in areas with a northeast aspect, intermediate gradients and depths between 200 and 1,000 m where the Deep Western Boundary Current is forced towards the surface by the local topography. These are the areas where Blainvilles beaked whales preferentially occurred.Communicated by O. Kinne, Oldendorf/Luhe     

Further insight on carapid—holothuroid relationships

Carapidae (or pearlfish) are eel-like fishes that live inside different invertebrates, such as holothurians, sea stars, or bivalves. Those of the genus Carapus are commensal and use their host as a shelter, while Encheliophis species are parasitic and eat the hosts gonads. In areas where they live in sympatry, C. boraborensis, C. homei, C. mourlani and E. gracilis are able to inhabit the same host species. Infestation is considered as monospecific when several conspecifics are observed in the same host. However, many aspects of this particular relation remain obscure, e.g. communication between carapids and the defence systems of the different protagonists (carapids and hosts). Experiments have been conducted in the field and laboratory to investigate several aspects of the carapids relationships with their hosts. Sampling carried out in the Bay of Opunohu (Moorea, French Polynesia) determined the sex ratio of C. boraborensis (3:1) and C. homei (1:1) and their distribution rate within different Echinodermata. Our study showed that neither species was capable of determining whether a heterospecific already occupied a sea cucumber or not. They were, however, able to locate the sea cucumbers cloaca, due to the excurrent resulting from respiration. The sea cucumbers defence system (Cuverian tubules) minimises predator attacks, but is not effective against carapid intrusion. The Carapidae defence system is twofold. Due to a passive system related to the sea cucumbers low cloacal position, the Cuverian tubules are not expelled when fish enter the cloaca. Moreover, carapids resist sea cucumber toxins better than other reef fish. Their increased resistance might be related to their gills rather than to their mucus coating; however, the latter may assist the fish in resisting the sticky substances emitted by the Cuverian tubules.Communicated by S.A. Poulet, Roscoff     

Rapid response to changing light environments of the calanoid copepod<Emphasis Type="Italic"> Calanus sinicus</Emphasis>

Calanus sinicus is a large calanoid copepod and a dominant species in the coastal waters of Japan. During a research cruise in Sagami Bay on 18 June 1996, we found C. sinicus performing an unusual diel vertical migration (DVM), a behavior that has not been reported in previous studies on this species. This study examined the DVM of C. sinicus under different light environments and revealed the copepods characteristic response to light. Field and laboratory results show that the DVM of C. sinicus is flexible and also confirmed its sensitivity and its rapid response to changing light environments. It is suggested that C. sinicus reacts to changes in absolute light intensity. This feature may be common in oceanic copepod species. The copepods quick reaction to light variation provides decreased predation risks and increased feeding opportunities, which make them a dominant survivor in coastal water habitats.Communicated by T. Ikeda, Hakodate     

Environmental influence on the locomotor activity rhythm of the sand-shore amphipod Talorchestia deshayesi

The precision and persistence of the nocturnal, surface locomotor activity rhythm of the sand-beach amphipod Talorchestia deshayesi is described. The rhythm is expressed as a circadian activity period with no evidence of a circa-tidal component associated with the time of high tide. The rhythm is light-sensitive and entrainment occurs in response to changes in the time of an experimentally simulated dusk transition. Locomotor activity is initiated at the end of the dusk transition with cessation always occurring prior to dawn. The point of activity initiation is taken to be the position of a synchronizing cue controlling entrainment. The cue appears to be an absolute irradiance level of approximately 1.0 to 2.0×10^-4 W m^-2     

Dufour’s gland pheromone as a reliable fertility signal among honeybee (Apis mellifera) workers

When a colony becomes queenless and without the possibility of requeening, honeybee workers initiate reproduction and lay male eggs about a week later. Assays in which two bees were confined in a small arena revealed that they establish a reproductive dominance hierarchy, i.e., one worker demonstrates greater ovarian development than her paired bee. Reproductive dominance is independent of relatedness, and can be established between full sisters, cousins, or random nestmates. A social environment, however, is obligatory, as singly housed bees fail to develop ovaries on the same time scale. Allowing varying degrees of social interactions between the paired bees revealed that olfaction of volatile bee compounds, as well as tactile communication, seem to provide the necessary social environment. Ovarian development was accompanied by the production of queen-like Dufours gland secretion in these workers. Especially notable was the increase in the queen-like esters. This increase was tightly linked to ovarian development and not necessarily to the dominance status of the bees in the pair. Thus, the occurrence of queen-like esters can serve as a reliable fertility signal. Advertising ovarian status may recruit helper workers with less developed ovaries (and which are less likely to successfully reproduce before colony breakdown) to assist their nestmates and thereby gain inclusive fitness. Revealing the role of Dufours gland secretion as a fertility signal adds another dimension to our understanding of how queen pheromones operate. The mandibular-gland secretion is a good predictor of dominance hierarchy, being correlated with false-queen characteristics but not fertility, whereas Dufours gland secretion is a good predictor of fertility but not dominance hierarchy.Communicated by R.F.A. Moritz     

Forage production and quality as influenced by amended quartz sand-tailings following phosphate mining

Quartz-sand tailings is a waste product from the Florida phosphate mining industry. Individual tailings disposal areas may occupy 20 to 60 ha and support no vegetation. A split plot field experiment was conducted on a sand tailings deposit to study the effect of various amendments on yield, quality, and mineral concentrations of Callie bermudagrass (Cynodon dactylon var.aridus Harlan et de Wet) and Siratro [Macroptilium atropurpureum (DC) Urb]. The two species were established on nine treatments: one sand-tailings control (SC); three with air-dried phosphatic clay (PC) at 110, 225, and 340 Mg ha^-1; and three with overburden (OB) at 448, 1,120, and 1,800 Mg ha^-1. All PC and OB treatment rates contained air-dried sewage sludge (SS) at 45 Mg ha^-1 and were all roto-tilled to a 20 to 25 cm depth. There were two additional treatments of OB at 1800 Mg ha^-1 without SS, with and without roto-tilling into sand tailings. Callie established slower (P<0.05) on SC and OB, than on treatments containing SS and/or PC. However, after the root system developed, yields were similar on all sand-tailings treatments. Crude protein andin vitro organic matter digestion for Callie and Siratro did not differ between soil treatments, but Siratro was generally superior to Callie in forage quality. Forage concentrations of P, K, Ca. Mg, Mn, and Zn were adequate in both species for good cattle growth on all treatments. However, Cu tended to be low in Siratro for plant and cattle growth. Once root systems were well established, good yields of quality forage with suitable mineral concentrations were produced on regularly fertilized, quartz-sand tailings.Contribution of the University of Florida Agric. Res. and Ed. Ctr., Ona, FL 33865, and Soil Sci. Dept., Univ. of Florida, Gainesville, FL 32611 as Journal Series No. 5575.     

Effect of light and turbulent mixing on the growth of<Emphasis Type="Italic"> Skeletonema costatum</Emphasis> (Bacillariophyceae)

Skeletonema costatum was grown in an outdoor mesocosm to test the hypothesis that fluctuations in irradiance brought about by changes in mixing time and depth can reduce diatom growth and biomass in the turbulent mixed layer. The light environment and mixing regime within the mesocosm were comparable to those in shallow lakes and coastal waters. Experiments showed no significant differences for 24-h mean and 7-day mean chlorophyll a and carbon-specific growth for mixed depths of 1 m and 3 m, and mixing times between 4 min and 65 min. Fluctuations in irradiance brought about by turbulent mixing had no significant effect on specific growth. The relationship between mixed depths and critical depths for S. costatum was therefore independent of fluctuations in irradiance in the turbulent mixed layer. The results indicated that to control growth of S. costatum mixed depths would have to exceed photic depths by a factor of 15, instead of the conventionally accepted factor of 5. Thus, it is likely that artificial mixing of shallow (<10 m) eutrophic waters will be more effective in controlling slow-growing summer biomass than fast-growing spring blooms dominated by diatoms.Communicated by P.W. Sammarco, Chauvin